ii Vol. 104 WINTER-SPRING 1996 Nos. 1-2 QL £/0T Journal of the New York Entomological Society (ISSN 0028-7199) Devoted to Entomology in General JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Editor: . Jame^ M. Carpenter, Department of Entomology, American Mu- seum of Natural History, Central Park West at 79th Street, New York, New York 10024 Book Review Editor: James S. Miller, Department of Entomology, Amer- ican Museum of Natural History, Central Park West at 79th Street, New York, New York 10024 Publications Committee: James K. Liebherr, Cornell University, Ithaca, Chairman; David L. Wagner, University of Connecticut, Storrs; Alfred G. Wheeler, Jr., Pennsylvania State Department of Agriculture, Harris- burg. 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Postmaster: Send address changes to the New York Entomological Society, 7o American Museum of Natural History, Central Park West at 79th Street, New York, New York 1(X)24-5192. Known office of publication. American Museum of Natural History, New York, New York 10024. Journal of the New York Entomological Society, total copies printed 700, paid circulation 602, mail subscription 602, free distribution by mail 19, total distribution 621, 79 copies left over each quarter. 0 This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). J. New York Entomol. Soc. 104(1-2): 1-20, 1996 THE SOUTH AMERICAN WEEVIL GENl (COLEOPTERA: CURCULIONIDAE; CRYPTORHYNCHINAE) Juan J. Morrone Laboratorio de Sistematica y Biologia Evolutiva (LASBE), Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina Abstract. — The genus Rhyephenes Schoenherr, endemic to the Central Chilean and Subant- arctic biogeographic provinces of southern South America, is easily recognized by its black body with shape resembling a spider. This genus includes seven species: Rhyephenes clathratus R. Philippi, R. gayi (Guerin), R. goureaui (Gay & Sober), R. humeralis (Guerin), R. lateralis (Guerin), R. maillei (Gay & Sober), and R. squamiger F. Philippi (reinstated herein from syn- onymy with R. gayi). A key, redescriptions, habitus photographs, and illustrations of the species are provided, and their geographical distribution is mapped. A cladistic analysis using 21 char- acters from external morphology and male and female genitalia produced four cladograms (Cl = 0.55, RI = 0.51, length 43 steps), which after successive weighting were reduced to one cladogram (Cl = 0.82, RI = 0.85, length 111 steps). In the cladogram, the following phylo- genetic sequence results: {R. squamiger, (R. lateralis, ((/?. clathratus, R. goureaui), (R. gayi, (R. humeralis, R. maillei))))). Biogeographic patterns exhibited by these species indicate a sequence from central Chile to southern Chile and Argentina. The weevil genus Rhyephenes (Curculionidae: Cryptorhynchinae) was described by Schoenherr (1837). This taxon of uncertain placement is one of the most char- acteristic weevil genera from central Chile, with two species extending also to south- ern Chile and Argentina. They are popularly known in Spanish as “aranitas” (little spiders), because of their typical habitus (see Figs. 1-14). Sixteen species-group names proposed by several authors (Guerin, 1830; Erichson, 1834; Gyllenhal, 1837; Gay & Sober, 1839; Blanchard, 1853; R. Philippi, 1859; E Philippi, 1899; Fiedler, 1942) were assigned to Rhyephenes. This nomenclatural proliferation is mainly due to the intraspecific variability of the species of Rhyephenes, which led authors to describe as valid species mere geographical variants. Recent synonymies by Kuschel (1993) and Morrone (1994) reduced the number of species to six. I suspected, how- ever, that the synonymy of R. squamiger E Philippi and R. gayi (Guerin) by Kuschel (1993) was incorrect. A cladistic analysis of its species can help ellucidate which are the valid species of Rhyephenes. MATERIAL AND METHODS The specimens examined in this study are from the following collections: AMNH, American Museum of Natural History, New York, USA (Lee Herman); BMNH, The Natural History Museum, London, Great Britain (Christopher Lyal); CWOB, Charles W. O’Brien private collection, Tallahassee, USA (Charles O’Brien); lADIZA, Insti- tute de Investigaciones de las Zonas Aridas, Mendoza, Argentina (Sergio Roig-Ju- nent); IPCN, Institute Patagonico de Ciencias Naturales, San Martin de los Andes, Argentina (Mario Gentili); MACN, Museo Argentine de Ciencias Naturales Bernar- dino Rivadavia, Buenos Aires, Argentina (Axel Bachmann); MHNS, Museo Na- 2 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Table 1. Data matrix and list of characters used in the cladistic analysis of Rhyephenes. 0 = plesiomorphic; 1, 2, 3 = apomorphic. outgroup 00000 00000 00000 00000 0 R. clathratus 11001 11001 21100 00010 0 R. gayi 10010 11000 21111 moo 1 R. goureaui 00001 11001 01101 OHIO 1 R. humeralis 21110 mil 21101 12101 1 R. lateralis 22000 10001 31000 01001 1 R. maillei 21010 mil 01000 moo 1 R. squamiger 00010 00101 10000 00100 0 1. Tubercles on male rostrum: (0) absent; (1) present, at sides; (2) present, on dorsum and sides. 2. Punctures on female rostrum: (0) broad, abundant; (1) fine, less abundant; (2) fine, sparse. 3. Prothorax: (0) slightly globose; (1) strongly globose, protuberant. 4. Relative width of the prothorax; (0) as wide as elytra; (1) wider than elytra. 5. Prothoracic anterior impression: (0) absent; (1) present. 6. Prothoracic punctures: (0) small, shallow; (1) large, deep. 7. Elytral basal area: (0) slightly impressed; (1) strongly impressed. 8. Elytral tubercles: (0) rounded; (1) subconical. 9. Elytral tubercles on intervals: (0) small, similar to those on striae; (1) large, more developed than those on striae. 10. Elytral tubercles on intervals and striae: (0) clearly separated; (1) partially fused. 11. Elytral humeral stripes of white scales: (0) absent; (1) two short stripes; (2) three short stripes; (3) three stripes, two short and one longer. 12. Body vestiture: (0) clothed with scales; (1) lacking scales. 13. Aedeagus in lateral view: (0) slender; (1) robust. 14. Ostiolar sclerites of aedeagus: (0) rounded; (1) acute. 15. Parameres: (0) long, narrow; (1) short, broad. 16. Female sternum 8 plate: (0) as long as wide; (1) longer than wide. 17. Relative length of female sternum 8 plate: (0) < 0.40X length of manubrium; (1) 0.41- 0.61 X length of manubrium; (2) > 0.6 IX length of manubrium. 18. Setae on sides of female sternum 8 plate: (0) absent; (1) present. 19. Arms of female sternum 8 plate: (0) narrow; (1) broad. 20. Spermathecal nodulus: (0) developed; (1) not developed. 21. Spermathecal ramus: (0) developed; (1) not developed. cional de Historia Natural, Santiago, Chile (Mario Elgueta); and MLP, Museo de La Plata, La Plata, Argentina (Juan Schnack). Drawings were made with a camera lucida attached to a stereoscopic microscope. Full data of type specimens are cited enclosing information from each label with square brackets with each line separated by a slash. Characters were derived from the external morphology, and male and female gen- italia. The classification of the subfamily Cryptorhynchinae is in some disarray (Lyal, 1993) and there are no keys available to the South American genera (O’Brien, 1984). Furthermore, the distinctness of Rhyephenes makes it presently not possible to iden- tify its sister group, so an unrooted ingroup analysis was performed, and the root of the cladogram was determined a posteriori using an hypothetical outgroup, following 1996 THE WEEVIL GENUS RHYEPHENES 3 the procedure described by Nixon and Carpenter (1993). In order to construct this outgroup, several South American genera were examined and the revision of the New Zealand Cryptorhynchinae (Lyal, 1993) was consulted. Table 1 contains the data matrix and the list of characters analyzed (all multistate characters were treated as additive). Analysis was carried out with Hennig86 1.5 (Farris, 1988), applying the implicit enumeration option and the successive weighting procedure. Cl and RI were calculated excluding autapomorphies. CLADOS 1.1 (Nixon, 1992) was em- ployed for examination of character distributions and for rooting the cladogram after the unrooted analysis. RHYEPHENES SCHOENHERR, 1837 Rhyephenes Schoenherr, 1837:312 (type species R. incas Gyllenhal [=R. humeralis (Guerin)], by original designation); Gay and Solier, 1839:24 (key); F. Philippi, 1899:3 (revision); Fiedler, 1942:274 (key). Physothorus Gay and Solier, 1839:22 (type species P. maillei Gay and Solier, by original designation); Solier, 1839:L {= Rhyephenes). Rhyephenes is easily recognized by its black color and habitus, which resembles a spider (Figs. 1-14). Biology: Adult Rhyephenes are usually seen on woody plants, females oviposit un- der bark of the host plants, and larvae are endophytic, excavating galleries and feeding on live wood (Angulo, 1970; Elgueta, 1993). Table 2 includes a list of the known plants where species of Rhyephenes have been collected, from personal ob- servations in the field, specimen labels, and relevant literature (Fiedler, 1942; Hav- rylenko & Winterhalter, 1949; Solervicens & Elgueta, 1989; Barriga et al., 1993; Elgueta, 1993; Morrone & Roig-Junent, 1995). These plants cannot be certainly assumed to be host plants. Distribution: The species of Rhyephenes occur in the Central Chilean and Subant- arctic provinces of the Andean subregion of southern South America (Figs. 50-53). KEY TO SPECIES OF RHYEPHENES 1. Elytra with 2-3 humeral stripes of white scales (Figs. 1, 3, 7, 9, 13) 2 Elytra lacking humeral stripes of white scales (Figs. 5, 11) 6 2. Prothorax strongly globose, protuberant (Fig. 7); elytra with tubercles on intervals more developed than those on striae R. humeralis (Figs. 7-8) Prothorax slightly globose; elytra with tubercles on intervals similar to those on striae 3 3. Elytra with basal area slightly impressed; aedeagus slender in lateral view (Figs. 24, 28) 4 Elytra with basal area strongly impressed; aedeagus robust in lateral view (Figs. 16, 18) 5 4. Body clothed with scales (Fig. 13); male rostrum lacking tubercles; female rostrum with broad, abundant punctures; prothorax wider than elytra, and with small, shallow punc- tures; elytra with subconical tubercles and two short humeral stripes (Fig. 13) R. squamiger (Figs. 13-14) Body lacking scales (Fig. 9); male rostrum with tubercles; female rostrum with fine, sparse punctures; prothorax as wide as elytra, and with large, deep punctures; elytra 4 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Figs. 1-8. Rhyephenes spp., habitus. 1, 3, 5, 7, dorsal view; 2, 4, 6, 8, lateral view. 1, 2, R. clathratus; 3, 4, R. gayi\ 5, 6, R. goureaui\ 7, 8, /?. humeralis. 1996 THE WEEVIL GENUS RHYEPHENES 5 Figs. 9-14. Rhyephenes spp., habitus. 9, 11, 13, dorsal view; 10, 12, 14, lateral view. 9, 10, R. lateralis', 11, 12, R. maillei', 13, 14, R. squamiger. with rounded tubercles and two short and one longer humeral stripes (Fig. 9) R. lateralis (Figs. 9-10) 5. Prothorax as wide as elytra and with anterior impression; elytra with tubercles partially fused; female rostrum with fine, less abundant punctures R. clathratus (Figs. 1-2) Prothorax wider than elytra and lacking anterior impression; elytra with tubercles clearly separated; female rostrum with broad, abundant punctures R. gayi (Figs. 3-4) 6. Prothorax wider than elytra and lacking anterior impression; elytra with basal area 6 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) strongly impressed, and subconical tubercles; aedeagus slender in lateral view; male rostrum with tubercles R. maillei (Figs. 11-12) Prothorax as wide as elytra and with anterior impression; elytra with basal area slightly impressed, and rounded tubercles; aedeagus robust in lateral view; male rostrum lacking tubercles R. goureaui (Figs. 5-6) Rhyephenes clathratus R. Philippi, 1859 (Figs. 1, 2, 15, 16, 29, 36, 43, 50) Rhyephenes clathratus R. Philippi, 1859:666; Fairmaire, 1860:251 {=R. goureaui)', F. Philippi, 1899:84 (reinstated). Rhyephenes clathratus was considered a synonym of R. goureaui by Fairmaire (1860). According to this analysis, they are sister taxa; the latter is distinguished by the elytra lacking the humeral stripes of white scales and by characters of the female genitalia. Redescription: Habitus (Figs. 1, 2). Body clothed with scales. Prothorax slightly globose, as wide as elytra; disc with anterior impression, and large and deep punc- tures. Elytra with basal area strongly impressed; with rounded tubercles; tubercles on intervals small, similar to those on striae, and partially fused to them; humeri with three short stripes of white scales. Male. Rostrum with tubercles at sides. Ae- deagus (Figs. 15, 16) robust in lateral view; ostiolar sclerites rounded; parameres long, narrow. Female. Rostrum with fine punctures. Sternum 8 (Fig. 29) with plate as long as wide, and sides lacking setae; arms broad. Hemistemite (Fig. 36). Sper- matheca (Fig. 43) with nodulus and ramus developed. Length 7.8-13.7 mm. Type material: Lectotype male (here designated: [UECTOFYV^IRhyephenesI clathratus/R. Philippi] [Rhyephenes/ clathratus/R. Phil./det. G. Kuschel/1990] (MHNS). Other material examined: CHILE. Without more precise data: 23 (1 AMNH, 3 BMNH, 19 MHNS). Cautm: Cautm, L. E. Pena, 1 (CWOB). Curico: La Montana, 8-IX-1970, A. Eglitis, 3 (MHNS). Limari: Las Trancas, 12-1-1978, D. Jackson, 1 (MHNS). Malleco: Malleco, XII- 1986, G. Perez de Arce, 1 (MHNS). Talca: Con- stitucion, pino insigne, 25-IX-1970, E. Holsten, 2 (MHNS); Los Cipreses, 1,000 m, 14-1-1968, L. E. Pena, 2 (MHNS). Rhyephenes gayi (Guerin, 1830) (Figs. 3, 4, 17, 18, 30, 37, 44, 50) Ty lodes gayi Guerin, 1830:125. Rhyephenes cacicus Gyllenhal, 1837:315. Rhyephenes gayi', Schoenherr, 1844:402 {=R. cacicus)', Lacordaire, 1866:108 (rein- stated). This species occurs abundantly in central Chile. It is identified by the acute ostiolar sclerite of the aedeagus. Redescription: Habitus (Figs. 3, 4). Body clothed with scales. Prothorax slightly globose, wider than elytra; disc lacking anterior impression, and large and deep punctures. Elytra with basal area strongly impressed; with rounded tubercles; tuber- 1996 THE WEEVIL GENUS RHYEPHENES 1 Table 2. Known plant associations of the species of Rhyephenes (plant families and species arranged alphabetically). -Asteraceae Ambrosia sp. R. gayi Bahia ambrosioides R. squamiger Baccharis concava R. squamiger Baccharis rhomboidalis R. squamiger Baccharis sp. R. gayi and R. humeralis -Berberidaceae Berberis rotundifolia R. humeralis -Boraginaceae Heliotropium stenophyllum R. squamiger -Bromeliaceae Puya chilensis R. squamiger -Celastraceae Maytenus boaria R. maillei -Fabaceae Adesmia microphylla R. gayi Anarthrophyllum sp. R. gayi Lupinus sp. R. humeralis Senna cummin gii R. gayi Senna multiglandulosa R. squamiger Senna sp. R. gayi -Juglandaceae Juglans regia R. humeralis -Lauraceae Persea americana R. squamiger -N othofagaceae Nothofagus antarctica R. maillei N. dombeyi R. humeralis and R. maillei N. nitida R. maillei N. pumilio R. maillei Nothofagus sp. R. humeralis and R. maillei -Pinaceae Pinus radiata R. clathratus, R. humeralis, and R. maillei -Rosaceae Prunus persica R. humeralis Quillaja saponaria R. maillei cles on intervals small, similar to those on striae, and clearly separated from them; humeri with three short stripes of white scales. Male. Rostrum with tubercles at sides. Aedeagus (Figs. 17, 18) robust in lateral view; ostiolar sclerites acute; para- meres short, broad. Female. Rostrum with broad, abundant punctures. Sternum 8 (Fig. 30) with plate longer than wide, and sides with setae; arms narrow. Hemistemite (Fig. 37). Spermatheca (Fig. 44) with nodulus developed and ramus not developed. Length 7.5-15.5 mm. Material examined: CHILE. Without more precise data: 102 (9 AMNH, 4 BMNH, 89 MHNS). Bio-Bio: Fundo Marfa Ester, 15 km W Victoria, 14-1-1989, M. Ramirez, 8 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Figs. 15-28. Rhyephenes spp., aedeagus. 15, 17, 19, 21, 23, 25, 27, dorsal view; 16, 18, 20, 22, 24, 26, 28, lateral view. 15, 16, R. clathratus; 17, 18, R. gayi\ 19, 20, R. goureaui\ 21, 22, R. humeralis\ 23, 24, R. lateralis', 25, 26, R. maillei', 27, 28, R. squamiger. 2 (MACN). Cachapoal: Termas de Cauquenes, 11-1-1953, 2 (MHNS). Cauquenes: Fundo El Roble, Colhueco, E Chilian, XII-1965, 24 (AMNH); Pelluhue, 31-VII- 1985, E Silva, 5 (MHNS), 18/20-IX-1985, E Silva, 1 (MHNS). Cautm: Cherquenco, 1954, 1 (MHNS). Chacabuco: Caleu, ex Salix vitiminalis, 10-III-1990, M. Elgueta, 1 (MHNS). Chiloe: Aucar, 6/15-1-1952, L. E. Pena, 2 (MHNS), 15-1-1952, L. E. Pena, 20 (MHNS). Choapa: Cerro La Silla del Gobemador, 31-X-1988. E. Maury, 1 (MLP); km 272 Panam. Norte, ex Senna cummingii var. coquimbensis, 14- V- 1985, M. Elgueta, 4 (MHNS), 23-1-1986, M. Elgueta, 1 (MHNS), exAdesmia microphylla, 11-III-1986, M. Elgueta, 1 (MHNS); Quilimari, 13-IX-1972, 1 (MHNS). Colchagua: Valle del Nilahue, 1-II-1915, 2 (MHNS). Concepcion: Concepcion, 26/28-XII-1926, E & M. Edwards, 1 (BMNH), 20-111-1952, 1 (MHNS). Cordillera: El Alfalfal, 23-11- 1967, V. Perez, 4 (MHNS), 25-1-1968; J. Moroni, 3 (MHNS). Curico: El Coigal, XI- 1955, L. E. Pena, 5 (CWOB). Elqui: La Marquesa, 350 m, A. Aguilera coll.. 1996 THE WEEVIL GENUS RHYEPHENES 9 Eigs. 29-49. Rhyephenes spp., female genitalia. 29-35, sternum 8, ventral view; 36—42, hemistemite, ventral view; 43-49, spermatheca. 29, 36, 43, R. clathratus', 30, 37, 44, R. gayi\ 31, 38, 45, R. goureaui; 32, 39, 46, R. humeralis; 33, 40, 47, R. lateralis-, 34, 41, 48, R. maillev, 35, 42, 49, R. squamiger. VIII-1973, 4 (MHNS); Pta. Lengua de Vaca, 31-VII-1971, L. E. Pena, 1 (MHNS). Los Andes: Estero Leiva, X-1953, 5 (MHNS); Rio Blanco, 30-XII-1917, 2 (MHNS). Malleco: Angol, 8-IX-1949, Cerda, 6 (MHNS), 18-III-1953, 1 (MHNS); Curacautm, 13-III-1952, 2 (MHNS). Melipilla: Cuesta de Barriga, 7-1-1952, 1 (MHNS); El Ca- nelo, 30-XI-1970, G. Bania, 1 (MHNS). Petorca: Quebrada del Chivato, 30-X-1988, E. Maury, 1 (MLP); Quebrada del Talanquen, 9-XII-1984, H. Niemeyer, 3 (MHNS); Quebrada El Tigre, 8-X-1988, E. Maury, 5 (MLP). Quillota: Palmas de Ocoa, 27-X- 1988, E. Maury, 1 (MLP); Quebrada Escobares, 22-11-1964, J. Solervicens, 3 (MHNS); Quillota, M. Fritz, 1 (MACN). San Felipe de Aconcagua: La Ollita, Cantillana, 2000 m, 18-XII-1969, L. E. Pena, 1 (MHNS). Santiago: Farellones, 2500 10 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) m, 4-XII-1969, J. Valencia, 1 (CWOB), 26-III-1971, J. Valencia, 2 (MHNS); Oasis, 11-1966, R. Perez, 17 (MHNS); Piche Alhue, 1800 m, 18-XII-1969, L. E. Pena, 3 (MHNS); San Cristobal, 14-XII-1966, R. Perez, 1 (MHNS); San Jose de Maipo, 17-11-1970, L. Alfaro, 1 (MHNS). Talagante: Talagante, 7-IX-1970, M. Elgueta, 4 (MHNS). Talca: Vilches Alto, Cord, de Talca, 1-1990, Arce, 5 (MHNS). Valdivia: La Union, Cocule, 6-III-1952, 2 (MHNS). Valparaiso: Marga Marga, 26-IV-1983, A. Mann, 2 (MHNS), 3 (BMNH); Quillota, sobre mspero, 1 (MHNS); Quintero, “bajo Ambrosia'\ 29-VIII-1981, 1 (MHNS); Valparaiso, XII-1925, A. Faz, 3 (BMNH). Rhyephenes goureaui (Gay and Solier, 1839) (Figs. 5, 6, 19, 20, 31, 38, 45, 51) Physothorus goureaui Gay and Solier, 1839:26. Rhyephenes goureaui', Solier, 1839:L. This species could be confused with R. maillei, the other species lacking humeral stripes in the elytra; the latter has the prothorax wider than the elytra, and the elytral tubercles on intervals more developed than those on striae. Redescription: Habitus (Figs. 5, 6). Body clothed with scales. Prothorax slightly globose, as wide as elytra; disc with anterior impression, and large and deep punc- tures. Elytra with basal area strongly impressed; with rounded tubercles; tubercles on intervals small, similar to those on striae, and partially fused to them; humeri lacking stripes of white scales. Male. Rostrum lacking tubercles. Aedeagus (Figs. 19, 20) robust in lateral view; ostiolar sclerites rounded; parameres short, broad. Female. Rostrum with broad, abundant punctures. Sternum 8 (Fig. 31) with plate as long as wide, and sides with setae; arms broad. Hemistemite (Fig. 38). Spermatheca (Fig. 45) with nodulus developed and ramus not developed. Length 8.0-15.2 mm. Material examined: ARGENTINA. Chubut: El Maiten, 28-XII-1958, A. Kovacs, 14 (AMNH). Neuquen: Catan-Lil, 25-11-1963, 1 (lADIZA); Lago Alumine, 25-11- 1964, 4 (lADIZA), 14-1-1993, G. Debandi, 1 (lADIZA); La Victoria, X-1963, 1 (lADIZA); without more precise data, 1 (MLP). CHILE. Without more precise data: 48 (1 AMNH, 2 MACN, 43 MHNS, 2 MLP). Arauco: P. N. Contulmo, 2-XI-1904, 1 (CWOB). Bio-Bio: Abanico, 19-1-1979, M. Elgueta, 1 (MHNS); San Carlos de Puren, 22-1-1973, 1 (MHNS). Cautm: Cherquenco, 1954, L. E. Pena, 3 (MHNS). Chiloe: Ancud, lO-IV-1914, 1 (AMNH), 19-XII-1926, F. & M. Edwards, 1 (BMNH); Pivehue, 16-11-1992, S. Roig-Junent, 1 (lADIZA). Curico: Potrero Grande, 23-X- 1976, G. Arriagada, 1 (MHNS). Llanquihue: Puerto Montt, 3 (MLP). Malleco: Cor- dillera Lonquimay, Sierra Nevada, 2-II-1962, J. Valencia, 1 (CWOB); P. N. Nahuel Buta, 1260 m, 23-XII-1985, S. Roig-Junent, 5 (lADIZA). Nuble: Cabreria, 31-1- 1977, F. Rodriguez, 1 (MHNS). Valdivia: Huellelhue, 24-1-1973, M. Elgueta, 1 (MHNS). Rhyephenes humeralis (Guerin, 1830) (Figs. 7, 8, 21, 22, 32, 39, 46, 52) Ty lodes humeralis Guerin, 1830:124. Cryptorhynchus arachnodes Erichson, 1834:264. 1996 THE WEEVIL GENUS RHYEPHENES 11 Rhyephenes incas Gyllenhal, 1837:314. Rhyephenes arachnodes; Gyllenhal, 1837:314 (=R. incas)\ Genuninger & Harold, 1871:2563 {=R. humeralis); Berg, 1899:154 (Luzon, erroneous type locality). Physothorus boyeri Gay and Solier, 1839:27. Rhyephenes boyeri; Solier, 1839:L (=R. incas); Lacordaire, 1866:108 (reinstated); Kuschel, 1955:287 {=R. humeralis). Rhyephenes humeralis; Schoenherr, 1844:402 {=R. incas); Gemminger & Harold, 1871:2563 (reinstated). Rhyephenes inca Lacordaire, 1866:108 (error noted by Berg, 1899:154). Rhyephenes aequalis F. Philippi, 1899:90; Kuschel, 1993:64 {—R. humeralis). This widespread species is variable in length. Its diagnostic strongly globose and protuberant prothorax can be less notable in some of the smaller specimens. Redescription: Habitus (Figs. 7, 8). Body clothed with scales. Prothorax strongly globose, protuberant, wider than elytra; disc lacking anterior impression, and large and deep punctures. Elytra with basal area strongly impressed; with subconical tu- bercles; tubercles on intervals large, more developed than to those on striae, and partially fused to them; humeri with three short stripes of white scales. Male. Ros- trum with tubercles on dorsum and sides. Aedeagus (Figs. 21, 22) robust in lateral view; ostiolar sclerites rounded; parameres short, broad. Female. Rostrum with fine punctures. Sternum 8 (Fig. 32) with plate longer than wide, and sides with setae; arms narrow. Hemistemite (Fig. 39). Spermatheca (Fig. 46) with nodulus and ramus not developed. Length 5. 5-6. 7 mm. Type material: Lectotype male (here designated): WJECTOTYPF.! Rhyephenes! aequalisIF. Philippi] [=Rhyepheneslhumeralisl(Gu6nn)l det. G. Kuschel/ 1990] (MHNS). Other material examined: ARGENTINA. Without more precise data: 10 (9 BMNH, 1 MACN). Mendoza: without more precise data, C. Bruch, 1 (MLP). Ne- uquen: without more precise data, 1 (MLP). Rio Negro: Bariloche, 16/18-XI-1926, F. & M. Edwards, 4 (BMNH). CHILE. Aisen: Puerto Cisnes, 11-1961, L. E. Pena, 6 (BMNH). Arauco: Arauco, 16-XII-1985, S. Roig-Junent, 2 (lADIZA), Baier, 5 (MACN); Temuco, 1-1975, S. Roig-Junent, 2 (lADIZA), 2 (MLP). Bio-Bio: Peme- hue, at night, 1-II-1968, L. & C. W. O’Brien, 6 (CWOB), 20-1-1993, G. Debandi, 1 (lADIZA). Cachapoal: Molloa, XI-I967, 1 (CWOB); Rengo, 11-1991, 4 (MLP). Car- denal Caro: Rosario, 23-11-1967, J. Numhauser, 2 (BMNH). Cautm: without more precise data, L. E. Pena, 5 (CWOB). Chacabuco: Caleu, ex Salix vitiminalis, lO-m-1990, M. Elgueta, 1 (MHNS). Chiloe: 25 km S Castro, V-1968, L. & C. W. O’Brien, 1 (CWOB); 9 km E Chepu, 4-II-1968, L. & C. W. O’Brien, 2 (CWOB). Choapa: 1 km S Huentelauquen, C. W. & L. O’Brien, 1 (CWOB); Los Vilos, 18-IX- 1969, A. Tobar, 13 (CWOB); Palo Colorado, N Quilimari, 27-X-1991, M. Elgueta, 2 (MHNS). Coihaique: Coihaique, 8-III-1972, A. Tobar, 2 (CWOB). Concepcion: Concepcion, 25-VIII-1968, J. Apablaza & C. W. O’Brien, 2 (CWOB), 21-III-1970, E. Holsten, 1 (MHNS), E. Reed, 1 (BMNH); 40 km E Concepcion, 25-VIII-1968, J. Apablaza & C. W. O’Brien, 1 (CWOB). Cordillera: San Jose de Maipo, 1-1969, R. Dulovitch, 1 (MLP). Curico: El Coigo, cordillera Curico, X-1964, 7 (AMNH), X-1974, 8 (AMNH); El Coiquel, XI-1955, L. E. Pena, 8 (CWOB); Quebrada Mala, 32 Km E Molina, 2-XI-1991, M. Elgueta, 1 (MHNS). Linares: 3 km SE Agua de 12 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) La Gloria, 4-XI-1967, 5 (CWOB); 13 km SE Agua de la Gloria, 4-XI-1967, L. & C. W. O’Brien, 4 (CWOB), 24-VIII-1968, L. & C. W O’Brien, 1 (CWOB); La Pasarela, 28 km de Linares, 1-1984, S. Roig-Junent, 2 (lADIZA); La Vega, 6 km S Parral, 8-XI-1991, M. Elgueta, 1 (MHNS); Panamavida, 20-X-1968, D. Correa, 2 (CWOB); Romehual, cordillera Parral, 5/10-XI-1960, L. E. Pena, 1 (IPCN). Los Andes: Los Andes, 22- V- 1979, Gordon, 2 (CWOB); Rio Blanco, 25-1-1974, L. E. Pena, 2 (MHNS). Malleco: Angol, 3-IX-1927, E. Reed, 1 (AMNH), 22-IV-1946, 2 (AMNH), 12-X-1946, S. Perez, 2 (AMNH), 20-VII-1947, W Mason, 2 (CWOB); 6 km W Angol, at night, 2-XI-1967, 5 (CWOB); cordillera Lonquimay, 4-1-1962, 1 (CWOB); R N. Nahuel Buta, 38 km W Angol, 4,300', 12-11-1968, L. & C. W O’Brien, 1 (CWOB); Pichinahuel, Cordillera Nahuel Buta, 1/10-1-1959, L. E. Pena, 1 (BMNH). Nuble: Fundo Las Cruces, Cordillera Parral, V-1958, L. E. Pena, 6 (BMNH), 12/14-X11-1960, L. E. Pena, 1 (BMNH); Las Trancas, 8 km W Termas de Chilian, 1300 m, 15-X1-1981, R. T. Schuh & N. 1. Platnick, 1 (AMNH); 15 km SE Recinto, at night, 1 -XI- 1967, L. & C. W O’Brien, 3 (CWOB). Petorca: Cachagua, 19-1X-1971, M. Elgueta, 1 (MHNS); 7 km NE Papudo, 8-V111-1968, L. & C. W. O’Brien, 2 (CWOB); 13 km S Quinquimo, 12-X1-1967, L. & C. W. O’Brien, 1 (CWOB); 10 km SE Zapallar, 11-11-1967, L. & C. W. O’Brien, 15 (CWOB), 23-lX- 1967, C. W. O’Brien, 1 (CWOB). Quillota: Limache, Cerro de la Cruz, 1975, S. Roig-Junent, 1 (lADlZA); Cerro Macaya, 18-X1-1962, A. Tobar, 1 (CWOB); 19 km E Manzanar, 3-X1-1967, L. & C. W. O’Brien, 1 (CWOB). San Antonio: antes de Los Queries, 15-1-1984, S. Roig-Junent, 2 (lADlZA). San Felipe de Aconcagua: 15 km SE Llay-Llay, 18-1X-1967, L. & C. W. O’Brien, 3 (CWOB). Santiago: 15 km E Arrayan, at night, C. W. O’Brien, 2 (CWOB); Cerro Robles, 100', at night, 24-X11-1968, L. & C. W. O’Brien, 6 (CWOB), 6300', 24 X11-1967, L. & C. W. O’Brien, 1 (CWOB); Curacavi, 14-X-1967, L. & C. W. O’Brien, 1 (CWOB), 23-Xl- 1967, C. W. O’Brien, 1 (CWOB); El Canelo, 5-X-1963, J. Valencia, 2 (CWOB), 19-X-1963, J. Valencia, 4 (CWOB), 2-X1-1963, J. Valencia, 1 (CWOB), 23-111-1968, D. Correa, 2 (CWOB); El Manzano, 23-X11-1967, D. Correa, 4 (CWOB), 6-111-1968, D. Correa, 1 (CWOB), 4-X-1970, 1 (MHNS); 2 km E El Manzano, 1-11-1969, D. Correa, 1 (CWOB); El Salto, 20-X11-1967, D. Correa, 1 (CWOB); Farellones, 8,600', under dung, 25-X11-1968, L. & C. W. O’Brien, 7 (CWOB), 2500 m, 4-X11-1969, J. Valencia, 2 (CWOB); La Reina, 10/15-X1-1967, J. Barros, 1 (CWOB), 6-X11-1967, C. Reyes, 1 (CWOB); Las Condes, 6-111-1938, 4 (BMNH); Las Cruces, 3-111-1974, E. Arriagada, 10 (CWOB); 5 km S Melipilla, at night, 16-1X-1967, L. & C. W. O’Brien, 7 (CWOB); 11 km S Melipilla, 300', at night, 16-1X-1967, J. Apablaza & C. W. O’Brien, 5 (CWOB); 8 km W Padre Hurtado, 3-1X-1967, L. & C. W. O’Brien, 2 (CWOB); 15 km W Padre Hurtado, at night, 6-1X-1969, L. & C. W. O’Brien, 4 (CWOB); P. N. La Campana, 14-111-1985, C. Vivan, 1 (MHNS); Rocas Sto. Domin- go, on lupine, 23-X1-1968, L. & C. W. O’Brien, 103 (CWOB); 4 km W Rungue, 6-1X-1968, L. & C. W. O’Brien, 1 (CWOB). Talagante: Naltagua, 10-11-1981, J. E. Barriga, 7 (MLP). Talca: Altos de Vilches, 30-X-1969, J. Rozen & L. E. Pena, 2 (AMNH), 27-X11-1969, J. Valencia, 2 (CWOB), lO-X-1970, A. Tobar, 4 (CWOB), 13-X-1970, A. Tobar, 1 (CWOB), 1280 m, 10/1 l-Xl-1971, J. Valencia, 3 (CWOB), 1200 m, 5/7-X-1972, Coscaron, Pena & Wygodzinsky, 7 (AMNH); Constitucion, 22-V-1969, E. Holsten, 3 (AMNH), 13-V-1970, E. Holsten, 2 (MHNS); 7 km W 1996 THE WEEVIL GENUS RHYEPHENES 13 Molina, M968, C. W. O’Brien, 10 (CWOB), 20-1-1968, D. Correa, 1 (CWOB), l-II- 1968, D. Correa, 30 (CWOB), 25-11-1968, on Salix trunk, D. Correa, 6 (CWOB), 26-X11-1968, D. Correa, 6 (CWOB); Valle Empedrado, 6-X-1970, E. Holsten, 1 (AMNH); Vilches Alto, 14-X11-1973, 2 (MHNS), 21-X11-1973, 1 (MHNS). Valdivia: Valdivia, D. Sharp, 2 (BMNH). Valparaiso: Algarrobo, on Baccharis, 24-X1-1968, L. & C. W O’Brien, 1 (CWOB), lX-1969, 1 (MHNS); 15 km NW Casablanca, 1000', at night, 9-1X-1967, L. & C. W O’Brien, 1 (CWOB); El Bato, E lllapel, 25-X-1991, M. Elgueta, 4 (MHNS); El Bato, a farm east of lllapel, 19-X-1966, 1 (AMNH); La Canela, 29-X-1991, M. Elgueta, 1 (MHNS); Valparaiso, 1903, Craw- ford expedition, 1 (BMNH), Walker, 1 (BMNH), E. Reed, 2 (AMNH), 3 (BMNH); 19 km SE Villa Alemana, 1,000', at night, lX-1967, L. & C. W O’Brien, 3 (CWOB); Vina del Mar, 17-111-1938, 1 (BMNH); 13 km S Vina del Mar, 650', 3-1X-1967, J. Barros, 1 (CWOB). Without more precise data: Araucama, R. M. Middleton, 13 (BMNH); A. Tobar, 6 (CWOB); Reed, 3 (BMNH); C. Darwin, 1 (BMNH); 116 (2 AMNH, 19 BMNH, 21 MACN, 70 MHNS, 4 MLP). Rhyephenes lateralis (Guerin, 1830) (Figs. 9, 10, 23, 24, 33, 40, 47, 52) Ty lodes lateralis Guerin, 1830:126. Rhyephenes lateralis; Schoenherr, 1844:404. This rather scarce species is easily recognized by the two short and one longer elytral humeral stripes. Redescription: Habitus (Figs. 9, 10). Body clothed with scales. Prothorax slightly globose, as wide as elytra, wider than elytra; disc lacking anterior impression, and large and deep punctures. Elytra with basal area slightly impressed; with rounded tubercles; tubercles on intervals small, similar to those on striae, and partially fused to them; humeri with three stripes of white scales, two short and one longer. Male. Rostrum with tubercles on dorsum and sides. Aedeagus (Figs. 23, 24) slender in lateral view; ostiolar sclerites rounded; parameres long, narrow. Female. Rostrum with fine, sparse punctures. Sternum 8 (Fig. 33) with plate as long as wide, and sides lacking setae; arms narrow. Hemistemite (Fig. 40). Spermatheca (Fig. 47) with no- dulus and ramus not developed. Length 9.1-13.2 mm. Material examined: CHILE. Without more precise data: 3 (MHNS). Nuble: Cob- quecura, 13-1-1967, P. Ramirez, 1 (CWOB); Quirihue, 1-1979, C. Vidal, 3 (MHNS). Talca: Alto Vilches, 14-III-1971, L. E. Pena, 2 (MHNS), 2-XI-1973, 1 (MHNS), 21-XII-1973, 2 (MHNS); Constitucion: 27-11-1948, 2 (MHNS), 25-IX-1970, E. Hol- sten, 5 (AMNH). Valdivia: Panguipulli, 14-11-1966, 1 (MHNS). Rhyephenes maillei (Gay and Sober, 1839) (Figs. 11, 12, 25, 26, 34, 41, 48, 53) Physothorus maillei Gay and Sober, 1839:24. Physothorus laevirostris Gay and Sober, 1839:25 (probably synonym of P. maillei). Rhyephenes maillei; Sober, 1839:L; Angulo, 1970:313 (larva). Rhyephenes laevirostris; Sober, 1839:L; Lacordaire, 1866:108 (not synonym of R. maillei); Berg, 1899:153 (=/?. maillei); F. Philippi, 1899:82 {=R. maillei); Hus- tache, 1936:235 (reinstated); Morrone, 1994:95 (=R. maillei). 14 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Rhyephenes immaculatus Blanchard, 1853:251, pi. 14, Fig. 16 {=R. laevirostris); Berg, 1899:153 (=R. maillei); Hustache, 1936:235 (=R. laevirostris); Morrone, 1994:95 (=R. maillei). Rhyephenes sulcatus F. Philippi, 1899:86; Morrone, 1994:95 {=R. maillei). Rhyephenes philippii Fiedler, 1942:279; Kuschel, 1950:18 {=R. sulcatus); Morrone, 1994:95 (=R. maillei). This widespread species is variable in length. The names R. laevirostris, R. im- maculatus, R. sulcatus, and R. philippii have been applied to variants of this species. Redescription: Habitus (Figs. 11, 12). Body clothed with scales. Prothorax slightly globose, wider than elytra; disc lacking anterior impression, and large and deep punctures. Elytra with basal area strongly impressed; with subconical tubercles; tu- bercles on intervals large, more developed than to those on striae; humeri lacking stripes of white scales. Male. Rostrum with tubercles on dorsum and sides. Aedeagus (Figs. 25, 26) slender in lateral view; ostiolar sclerites rounded; parameres long, narrow. Female. Rostrum with fine punctures. Sternum 8 (Fig. 34) with plate longer than wide, and sides with setae; arms narrow. Hemistemite (Fig. 41). Spermatheca (Fig. 48) with nodulus developed and ramus not developed. Length 6.3-15.4 mm. Type material: Lectotype male (here designated): {W.CTOTY¥F.IRhyephenesl sulcatus/¥. Philippi] [Rhyephenes/maillei/varmni Gay & Solier/det. G. Kuschel/ 1990] (MHNS). Other material examined: ARGENTINA. Chubut: El Hoyo, 5-V-1959, A. Kovacs, 4 (BMNH), 18-X-1960, A. Kovacs, 1 (BMNH); Lago Verde, 560 m, 4-II-1983, M. & P. Gentili, 1 (IPCN); Rio Correntoso, 1 (MACN); without more precise data, 2 (MLP). Mendoza: without more precise data, 1 (AMNH). Neuquen: Alumine, 25-11- 1964, 1 (lADIZA), 11-1977, O. de Ferraris, 3 (IPCN), 8-1-1991, G. Debandi, 2 (lA- DIZA); Chapelco, 7-III-1964, 2 (lADIZA), 1,700 m, 15-III-1964, M. Gentili, 3 (IPCN), 1750 m, XII-1965, M. Gentili, 1 (IPCN), 1400 m, 30-III-1982, M. Gentili, 1 (IPCN); Copahue, 1,925 m, 18-XII-1963, M. Gentili, 1 (IPCN), 2000 m, 3-II-I987, M. Gentili, 1 (IPCN); Huiliches, 13-XII-I99I, G. Debandi, 1 (lADIZA); Kilca, 4-I-I972, 1 (IPCN); Lago Hermoso, XII-1949, M. Barrera, 3 (MACN); Lago Lacar, 23 km W San Martin de los Andes, 25-1-1972, L. Herman, 1 (CWOB); Lago Lacar, Nonthue, 640 m, 17-XI-1992, M. Gentili, 2 (IPCN); Lago Queni, 875 m, M. Gentili, 1 (IPCN); Lago Tromen, 1,000 m, 15-XII-1962, M. Gentili, 1 (IPCN); Pucara, 1949, S. Schajovskoi, 2 (MACN), 23-XI-1957, M. Gentili, 1 (IPCN), 21-XII-I965, L. Grosso, 3 (MLP), 6-II-1972, L. Herman, 1 (CWOB); Rio Alumine, 16-XII-1965, M. Gentili, 1 (IPCN); road between Pucara and Lago Venado, 24/25-1-1972, L. Herman, 2 (CWOB); San Martin de los Andes, 4-III-1942, M. Biraben, 1 (MLP), 17-III-1964, 1 (lADIZA), 9-XII-1976, O. de Ferraris, 1 (IPCN), 1000 m, 2-XI-1986, M. & P. Gentili, 1 (IPCN), 640 m, M. Gentili, 5 (IPCN), 10-XI-199I, G. Debandi, 2 (lA- DIZA); valle del Rio Limay, 11-1947, J. Navas, 3 (MLP); without more precise data, 28 (24 MACN, 4 MLP). Rio Negro: Bariloche, 25/28-X-1926, F. & M. Edwards, 2 (BMNH), 11-1938, Biraben & Scott, 1 (MLP), 1-1942, Rossi, 3 (MACN), 26-XI- 1963, A. Kovacs, 1 (AMNH), Richter, 5 (MLP); El Bolson, 20-IX-1956, A. Kovacs, 2 (BMNH), 12-VII-1958, A. Kovacs, 1 (BMNH), 25-XII-1958, A. Kovacs, 16 (BMNH), 18-XII-1959, A. Kovacs, 1 (BMNH), 22-XII-1959, A. Kovacs, 1 (BMNH), 29-XI-1959, A. Kovacs, 1 (BMNH), 5-X-1960, A. Kovacs, 6 (BMNH), 18-1-1961, 1996 THE WEEVIL GENUS RHYEPHENES 15 1 (BMNH), lO-VIII-1961, A. Kovacs, 1 (BMNH), 20-X-1961, 2 (BMNH), 1-XII- 1961, A. Kovacs, 4 (BMNH), 9-XIM962, A. Kovacs, 1 (BMNH), 22-XII-1962, A. Kovacs, 1 (BMNH), 16-1-1963, A. Kovacs, 1 (BMNH), 19-1-1965, A. Kovacs, 11 (AMNH); isla Victoria, 15-XI-1969, 1 (MACN), 2-XII-1969, 1 (MACN); isla Vic- toria, IV-1958, 2 (lADIZA); Norquinco, 10-11-1964, A. Kovacs, 1 (AMNH); R N. Los Alerces, S. Roig-Junent, 1 (lADIZA); Pto. Radal, isla Victoria, 1 (MACN). Santa Cruz: Lago Argentino, 1959, R W. James, 3 (BMNH), 16/27-11-1974, C. Bordon, 1 (CWOB); valle del Lago Blanco, 1903, 2 (BMNH); Valle Tunel, 3 (MLR). Tierra del Fuego: Without more precise data, 1900, C. Bruch, 6 (MACN). Without more precise data: Patagonia, 69 (MLR); 1935, 5 (BMNH), 1-1972, L. Herman, 1 (CWOB). CHILE. Without more precise data: 50 (5 AMNH, 12 BMNH, 5 MACN, 26 MHNS, 2 MLR). Aisen: Lago J. A. Rios, Taitao, 8-XII-1956, C. Riffart, 1 (MHNS); Pto. Aisen, 12-III-1943, R. Maldonado, 39 (MLR), 21-1-1968, L. & C. W. O’Brien, 1 (CWOB); Taitao, Base no. 5, 5-XII-1956, C. Bittart, 2 (MHNS); without more pre- cise data, 11-1934, 1 (MHNS). Arauco: Arauco, 24-III-1969, G. Billings, 7 (MHNS); Canete, 24-III-1969, R. Billings, 6 (MHNS), pino insigne, 12- V- 1969, D. Edelman, 1 (AMNH); Caramavida, 31-XII-1957, L. E. Pena, 1 (MHNS), 18-X-1969, G. Bama, 1 (MHNS), 16-XII-1985, S. Roig-Junent, 2 (lADIZA); Curanilahue, l-IV-1970, D. Edelman, 1 (AMNH); isla Mocha, XI- 1982, Bullock, 3 (MHNS); P. N. Contulmo, 19-X-1969, G. Barria, 1 (MHNS), 15-XII-1985, S. Roig-Junent, 5 (lADIZA); without more precise data, Baier, 1 (MACN). Bio-Bio: Mulchen, 16-11-1969, F. Rojas, 1 (MHNS). Capitan Pratt: Dos Lagunas, night, 13-1-1968, C. W. & L. O’Brien, 1 (CWOB), under wood, 14-1-1968, C. W. & L. O’Brien, 5 (CWOB). Cautin: rio Cautm, L. E. Pena, 2 (CWOB); 50 km S Temuco, ll-XII-1984, S. Roig-Junent, 1 (lADIZA); Villarica, 3-II-1968, B. Heineman, 1 (AMNH). Chiloe: Aucar, 18-1-1952, L. E. Pena, 7 (MHNS), 6-V-1952, L. E. Pena, 2 (MHNS); 9 km N Castro, 7-II-1968, L. & C. W. O’Brien, 1 (CWOB); 14 km E Chepu, 4-II-1968, L. & C. W. O’Brien, 3 (CWOB); Dalcahue, 18-1-1962, R, Usinger, 1 (CWOB); 20 km N Pichue, on [Nothofagus] nitida, 16-XI-1992, S. Roig-Junent, 1 (lADIZA); 6 km N Quellon, 6-II- 1968, L. & C. W. O’Brien, 1 (CWOB); 21 km N Quellon, 6-II-1968, L. & C. W. O’Brien, 1 (CWOB); without more precise data, C. Darwin, 1 (BMNH). Coihaique: Coihaique, 1-1934, 9 (MHNS); 17 km SE Coihaique, 20-1-1968, L. & C. W. O’Brien, 4 (CWOB); 56 km SE Coihaique, 20-1-1968, L. & C. W. O’Brien, 1 (CWOB); 64 km SE Coihaique, 20-1-1968, L. & C. W. O’Brien, 12 (CWOB); 7 km W Coihaique, at night, 21-1-1968, L. & C. W. O’Brien, 1 (CWOB); 33 km W Coihaique, 23-1- 1968, L. & C. W. O’Brien, 4 (CWOB). Coichagua: 10 km S San Fernando, 12-XI- 1967, J. Undurraga, 1 (CWOB). Concepcion: Concepcion, 21 -III- 1969, E. Holstem, 1 (AMNH). Llanquihue: El Manso, 22-1-1959, A. Kovacs, 13 (BMNH), l-IV-1960, A. Kovacs, 52 (BMNH); Frutillar Bajo, 11-1965, G. Silva, 2 (MHNS); Pto. Varas, M. Richter, 2 (MACN). Magallanes: Cabo Negro, 28-XII-1975, Perez, 6 (MHNS); cordillera Lonquimay, Sierra Nevada, 2-1-1962, J. Valencia, 1 (CWOB), 4-1-1962, 1 (CWOB); Dawson island, 1916, 1 (BMNH); El Canelo, 15-III-1969, L. E. Pena, 3 (MHNS); Estancia Canelo, 17-1-1968, L. & C. W. O’Brien, 12 (CWOB); 26 km E Estancia Canelo, on and under Nothofagus, 17-1-1968, L. & C. W. O’Brien, 37 (CWOB); isla Navarino, 1935, J. Bird, 1 (AMNH); Laguna Amarga, 10-1-1968, C. W. & L. O’Brien, 1 (CWOB); Magallanes, 2-IV-1953, T. Cekalovic, 1 (MHNS); Punta Arenas, Walker, 2 (BMNH); 3 km W Punta Arenas, at night, 16-1-1968, L. & 16 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) C. W. O’Brien, 1 (CWOB). Malleco: Icalma, 11-1989, 1 (MHNS); Pailahueque, X-1962, M. Fritz, 3 (MACN); R N. Nahuel Buta, 1200 m, 23-XII-1985, S. Roig- Junent, 3 (lADIZA), 20-1-1993, G. Debandi, 2 (lADIZA), 21-1-1993, G. Debandi, 1 (lADIZA); P. N. Tolhuaca, 13-1-1993, G. Debandi, 2 (lADIZA); Pichinahuel, Na- huelbuta, 1200 m, 14-11-1956, L. E. Pena, 1 (CWOB). Osomo: Antill^lnca, 3,700', Nothofagus pumilio, 10-11-1968, C. W. O’Brien, 1 (CWOB); 18 km NW Antillanca, 1,800', at night, 9-II-1968, L. & C. W O’Brien, 3 (CWOB); Bahia Mansa, 26-IX- 1968, C. Calderon, 3 (MHNS); Osomo, 1-1975, S. Roig-Junent, 9 (lADIZA), XII- 1984, S. Roig-Junent, 1 (MACN); 51 km E Osomo, Lago Puyehue, 800', 9-II-1968, L. & C. W. O’Brien, 8 (CWOB); R N. Puyehue, 11-1992, S. Roig-Junent, 1 (lADI- ZA); P. N. Puyehue, Agua Caliente, XII-1984, S. Roig-Junent, 1 (lADIZA); Puca- trihue, coastal town, 3/21 -III- 1967, 2 (AMNH); Purranque, IX- 1985, J. E. Barriga, 1 (MHNS). Santiago: Lago Arrayan, 20-XII-1968, C. Barrientos, 1 (CWOB). Tala- gante: Maipo, 7-XI-1968, J. Marshall, 1 (CWOB). Talca: Alto de Vilches, 10-X- 1970, A. Tobar, 1 (CWOB). Tierra del Fuego: forest S of Bahia Iniitil (Useless Bay), XII-1904, R. Crawshay, 1 (BMNH); no McKlelland, XI/XII-1904, R. Crawshay, 1 (BMNH). Ultima Esperanza: Cancha Carreras, 24-1-1988, J. J. Morrone, 1 (MLP); Cerro Guido, Nothofagus, 10-1-1968, C. W. & L. O’Brien, 4 (CWOB); Cerro Leon, 19-1-1969, V. Perez, 1 (MHNS); P. N. Torres del Paine, 25-1-1988, J. J. Morrone, 1 (MLP). Valdivia: Lanco, 1-1980, 4 (MLP); La Union, 10-III-1952, 1 (MHNS); Pan- guipuli, 14-11-1966, G. Ortega, 20 (CWOB), 11-1984, L. E. Pena, 1 (CWOB); Pire- hueico, l-XII-1985, S. Roig-Junent, 1 (lADIZA); Valdivia, 30-XII-1968, D. Correa, 1 (CWOB), 31-XII-1968, night, D. Correa, 9 (CWOB), 3-1-1969, D. Correa, 56 (CWOB), 29-XII-1986, E. Krammer, 2 (MHNS); 13 km SE Valdivia, 3-II-1968, L. & C. W. O’Brien, 1 (CWOB); 18 km N Valdivia, 4-1-1969, D. Correa, 11 (CWOB). Rhyephenes squamiger F. Philippi, 1899, reinstated name (Figs. 13, 14, 27, 28, 35, 42, 49, 53) Rhyephenes squamiger F. Philippi, 1899:88; Kuschel, 1993:63 {=R. gayi). This scarce species, easily recognized by its body clothed with scales, has a re- lictual distribution in central Chile. Kuschel (1993) incorrectly considers this species as a geographical race of R. gayi; however, the cladistic analysis indicates that they are two different (and unrelated) species. Redescription: Habitus (Figs. 13, 14). Body lacking scales. Prothorax slightly glo- bose, wider than elytra; disc lacking anterior impression, and small and shallow punctures. Elytra with basal area slightly impressed; with subconical tubercles; tu- bercles on intervals small, similar to those on striae, and partially fused to them; humeri with two short stripes of white scales. Male. Rostmm lacking tubercles. Aedeagus (Figs. 27, 28) slender in lateral view; ostiolar sclerites rounded; parameres long, narrow. Female. Rostmm with broad, abundant punctures. Sternum 8 (Fig. 35) with plate as long as wide, and sides with setae; arms narrow. Hemistemite (Fig. 42). Spermatheca (Fig. 49) with nodulus and ramus developed. Length 9.1-10.9 nun. Type material: Lectotype male (here designated): [883/a] [squamiger^ [= Rhyephe- nes/gay i/(Gu6nn)/det. G. Kuschel/1990] [LECTOTYFE/ Rhyephenes/squamiger/F. Philippi] (MHNS). Other material examined: CHILE. Choapa: Los Vilos, “ex Senna cummingV', 1996 THE WEEVIL GENUS RHYEPHENES 17 Figs. 50-53. Rhyephenes spp., geographical distribution. 50, R. clathratus (black circles) and R. gayi (open circles); 51, R. goureaui; 52, R. humeralis (black circles) and R. lateralis (open circles); 53, R. maillei (black circles) and R. squamiger (open circles). 18 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) — root 4 8 H — I — R. squatniger 1 6 11 12 17 21 111 I I I 112 111 1 2 11 18 20 \ — I — I — I — I — R. lateralis 2 2 3 0 1 5 19 ^-+ 7 13 4-+ 2 17 18 21 H-HHh R. clatbratus 10 0 0 1 11 15 R.goureaui 0 0 1 10 14 |— R.gayi 4 15 16 4-I-+ 0 1 12 8 9 t- f I I 2 111 3 17 20 t-H- R. humeralis Fig. 54. squares = 1 2 1 11 13 15 HUH- R maiUei 0 0 0 Cladogram of the species of Rhyephenes. Black squares = synapomorphies; dotted homoplasies. 11-1989, Barriga, 1 (MHNS); Nague, Los Vilos, 17-IX-1971, L. E. Pena, 1 (MHNS); km 27 Panamericana Norte, ex Bahia ambrosioides, 24-IX-1985, 1 (MHNS). Limarf: Fray Jorge, at night, 3-X-1967, L. & C. W. O’Brien, 1 (CWOB), 20- VI-1968, L. & C. W. O’Brien, 1 (CWOB), 21-VI-1968, L. & C. W. O’Brien, 2 (CWOB), 7-X-1975, C. Vidal, 1 (MHNS). CLADISTICS AND BIOGEOGRAPHY The analysis of the data matrix (Table 1) produced four cladograms (Cl = 0.55, RI = 0.51, length 43 steps) under equal weights, which after successive weighting resulted in one cladogram (Cl = 0.82, RI = 0.85, length 111 steps). In the cladogram (Fig. 54), the following phylogenetic sequence results: (R. squamiger, (R. lateralis, {{R. clathratus, R. goureaui), {R. gayi, (R. humeralis, R. maillei))))). When the cladogram is compared with the distributional maps of the species of Rhyephenes (Figs. 50-53), basal species {R. squamiger and R. lateralis) appear to be endemic to central Chile, whereas the most apomorphic species are widespread in central and southern Chile {R. gayi) or extend also to southern Argentina (R. humeralis and R. maillei). Whether this pattern reflects a series of vicariant or dis- persal events cannot be deduced directly from the cladogram sequence. Future com- parisons with other plant and animal taxa distributed in the same areas will help to solve this problem. AC KNO WLEDGMENTS I thank Adrian Fortino for inking the drawings and Federico Ocampo for taking the photo- graphs. This study was supported by grant 4662-91 of the National Geographic Society and the Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET), Argentina. 1996 THE WEEVIL GENUS RHYEPHENES 19 LITERATURE CITED Angulo, A. O. 1970. Descripcion de la larva y pupa de Rhyephenes maillei (Gay & Solier), (Coleoptera: Curculionidae). Bol. Soc. Biol. Concepcion, 1969-70, 42:313-316. Barriga, J. E., T. Curkovic, T. Fichet, J. L. Henriquez and J. Macaya. 1993. Nuevos antece- dentes de coleopteros xilofagos y plantas hospederas en Chile, con una recopilacion de citas previas. Rev. Chil. Entomol. 20:65-91. Berg, F. G. C. 1899. El genero Rhyephenes Schoenh. en la Republica Argentina. Comun. Mus. Nac. B. Aires 1(5): 15 1-154. Blanchard, C. E. 1853. Insectes. In: Voyage au Pole Sud et dans I’Oceanie sur les corvettes r Astrolabe et la Zelee; execute par ordre du Roi pendant les annees 1837-1838-1839- 1840 sous le commandement de M. J. Dumont-d’Urville, Capitaine de vaisseau. Zool., vol. 4, part 1, Baudry, Paris, 422 pp. Elgueta, M. 1993. Las especies de Curculionoidea (Insecta: Coleoptera) de interes agricola en Chile. Publ. Ocas. Mus. Nac. Hist. Nat., Chile 48:5-79. Erichson, W. E 1834. Coleoptera, pp. 219-276. In: Meyen, E J. E, Beitrage zur Zoologie, gesammelt auf einer Reise urn die Erde. Nova Acta Acad. Caes. -Leopold. -Carol. Nat. Cur. 16(Suppl. l):219-284. Fairmaire, L. 1860. in: Dohm, C. A., Nachschrift der Redaction. Stett. Entomol.Ztg. 21(7- 9):25 1-252. Farris, J. S. 1988. Hennig86 reference. Version 1.5. Published by the author. Port Jefferson, New York. Fiedler, C. 1942. Die Riisselkafergattung Rhyephenes Schonh. (Coleoptera: Curculionoidea: Cryptorhynchidae). Arb. Morphol. Taxon. Entomol. Berlin- Dahlem 9(4):274-279. Gay, C. and A. J. J. Solier. 1839. Memoire sur deux genres remarquables de Curculionites du Chili. Ann. Soc. Entomol. France 8:5-82. Gemminger, M. and E. von Harold. 1871. Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus. Vol. 8, Curculionidae. Gummi, Monachii, pp. 2181-2668. Guerin, E E. 1830. Crustaces, arachnides et insectes. in: Duperrey, L. L., Voyage autour autour du monde, execute par ordre du Roi, sur la corvette de Sa Majeste, La Coquille, pendant les annes 1822, 1823, 1824 et 1825, sous le Ministere et conformement aux instructions de S. E. M. le Marquis de Clermont-Tonnerre, Ministre de la Marine. Zoologie, vol. 2, part 2, div. 1, Bertrand, Paris, 319 pp. Gyllenhal, L. 1837. in: Schoenherr, C. J. (1837). Havrylenko, D. and J. J. Winterhalter. 1949. Insectos del Parque Nacional Nahuel Huapi. Minist. Obras. Servic. Publicos, Adm. Parques Nac. y Turismo, 47 pp. Hustache, A. A. 1936. Coleopterorum Catalogus. Junk, ’s-Gravenhage. Pars 151, Curculioni- dae: Cryptorhynchinae, pp. 1-137. Kuschel, G. 1950. Nuevas sinonimias, revalidaciones y combinaciones (9° aporte a Col. Cur- culionidae). Agric. Tec., Chile 10(1):10-21. Kuschel, G. 1955. Nuevas sinonimias y anotaciones sobre Curculionoidea (Coleoptera). Rev. Chil. Entomol. 4:261-312. Kuschel, G. 1993. in: Elgueta (1993). Lacordaire, J. T. 1866. Histoire naturelle des insectes. Genera des Coleopteres ou expose methodique et critique de tous les genres proposes jusqu’ici dans cet ordre d’ insectes. Vol. 7. Roret, Paris, 620 pp. Lyal, C. H. C. 1993. Cryptorhynchinae (Insecta: Coleoptera: Curculioninae). Fauna of New Zealand 29:1-302. Morrone, J. J. 1994. Nomenclatural changes in Patagonian Belidae and Curculionidae (Cole- optera: Curculionoidea). Neotropica 40(103-104):95. Morrone, J. J. and Roig-Junent. 1995. The diversity of Patagonian weevils: An illustrated 20 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) checklist of the Patagonian Curculionoidea (Insecta: Coleoptera). L.O.L.A., Buenos Ai- res, 189 pp. Nixon, K. C. 1992. CLADOS ver. 1.1. IBM PC-compatible character analysis program. Pub- lished by the author. Nixon, K. C. and J. M. Carpenter. 1993. On outgroups. Cladistics 9(4);413-426. O’Brien, C. W. 1984. Revision of the Neotropical weevil genus Philonis (Cryptorhynchinae: Curculionidae: Coleoptera). Southwest. Entomol. 9(2):232-239. Philippi, F. H. 1899. Monografia del jenero Rhyephenes Schoenh. An. Univ. Chile 104:81-93. Philippi, R. A. 1859. Algunas especies nuevas de Coleopteros de la provincia de Valdivia. An. Univ. Chile 16:656-678. Schoenherr, C. J. 1837. Genera et species curculionidum, cum synonymia hujus familiae. Roret, Paris. Vol. 4, part 1, 600 pp. Schoenherr, C. J. 1844. Genera et species curculionidum, cum synonymia hujus familiae. Roret, Paris. Vol. 8, part 1, 442 pp. Solervicens, J. and M. Elgueta. 1989. Entomofauna asociada al matorral costero del Norte Chico. Acta Ent. Chil. 15:91-122. Solier, A. J. J. 1839. Rectifications importantes a faire au memoire sur deux genres remarqu- ables de Curculionites du Chili (page 6 de ce vol.). Bull. Soc. Entomol. France 1839: XLIX-LI. Received 24 June 1996; accepted 27 February 1997. J. New York Entomol. Soc. 104(1— 2);21— 36, 1996 CHECKLIST OF THE SPECIES IN THE SUBFAMILY STENOGASTRINAE (HYMENOPTERA: VESPIDAE) James M. Carpenter and Jun-ichi Kojima Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA and Natural History Laboratory, Faculty of Science, Ibaraki University, Mito 310, Japan Abstract. — A checklist of the species in the subfamily Stenogastrinae is presented, including synonyms, nomenclatural changes, and distributional summaries. Forty-five species in seven genera are treated as valid, with an additional three subspecies. Excluding emendations, mis- spellings and nomina nuda, a further 16 names are treated as available synonyms, but eight of these names are listed only as questionable synonyms. A new combination is Parischnogaster aurifrons (Smith). Thirty new locality records are given. The taxonomy of the hover wasps, Stenogastrinae, has not been well studied, compared to to the other two social wasp subfamilies (Vespinae and Polistinae). Well corroborated taxonomic works would of course contribute greatly to the study of the biology of these wasps, which exhibit considerable divergence in social organization and nesting behavior (reviewed in Turillazzi, 1991). A check- list, which provides basic taxonomic information, such as the currently available species names, synonymy and type depositories, is helpful in perusing revisionary taxonomic works and catalogs. The checklist presented here gives such information for the Stenogastrinae. It covers all the nomenclatural changes and provides also a summary of distribu- tional data. The last attempt to list all of the Stenogastrinae was by von Schulthess (1927). Although useful, von Schulthess did overlook a few available names when compiling his list, and 26 taxa have since been described. This includes six genera. Five of these genera were described by the late J. van der Vecht (1969, 1972, 1977), who also described most of the species level taxa (1972, 1975, 1977). Van der Vecht was preparing taxonomic revisions of all the Sten- ogastrinae, but died before he published the revisions of Eustenogaster, Liosten- ogaster and Parischnogaster. The revisions van der Vecht began are now being carried on by C. K. Starr and S. Turillazzi, and a few descriptions have appeared (Turillazzi, 1988; Turillazzi and Carfi, 1996). In the meantime, over the course of his work van der Vecht distributed specimens labelled with no fewer than 24 manuscript names to various collections. One consequence of this was predict- able: two of these names have been published as nomina nuda (Hansell, 1987; Carpenter, 1988; see below). Van der Vecht’s unpublished notes also indicated a few synonymies that have still not appeared. These are listed below only as questionable synonyms, because we ourselves have not yet studied all of the relevant types. But two of these synonymies, if correct, entail the replacement of names currently in use (namely Parischnogaster alternata and P. timida). We are not taking the step of replacing these names, but are listing the putative senior 22 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) synonyms, in order to draw the attention of other scientists to the potential syn- onymy, and so encourage study of the possibility. Moreover, potential synonyms have been overlooked in regional works (Das and Gupta, 1984, 1989; Reyes, 1988), which are thus incomplete. Another peculiarity of this piecemeal publication of the taxonomy for the sub- family is that most of the species were transferred from the two genera recognized by von Schulthess (1927) to the modern genera in publications devoted to other subjects. This includes the cladistic analysis by Carpenter (1982, 1988), and the book on the ethology of Hymenoptera written by Iwata. The combinations ap- peared in Iwata’s book apparently because the wasps observed by Iwata were identified by van der Vecht. Although Iwata used these modern combinations in his original book written in Japanese (1971), the English translation published in 1976 is cited in the checklist because of the easier availability of the latter book, and because no nomenclatural changes were made for the species Iwata treated between 1971 and 1976 (except Krombein, 1976). The arrangement of the checklist is alphabetical. The status given each taxon generally follows the most recent author to treat that taxon. Each name is fol- lowed by the original citation, followed by synonymies and generic changes, which are listed chronologically. The unpublished synonyms in van der Vecht’s notes are listed with question marks preceding. Misspellings are indicated by exclamation points in brackets. Each publication is cited in abbreviated form. Where two dates are listed, the first is the true date of publication, so far as can be determined, with the date following in parentheses being the date printed on the publication. The original locality is quoted, and the type depository is then listed in parentheses, as the city containing the collection. Distributional data are summarized by country in the final paragraph for an taxon treated as valid. An asterisk denotes a new locality record. Phylogenetic relationships among the genera are: Liostenogaster + {{Eusten- ogaster + Stenogaster) + {Anischnogaster + (Metischnogaster + Parischnogas- ter))) (Carpenter, 1988). Carpenter (1988) synonymized one of van der Vecht’s (1977) genera, sinking Holischnogaster into Parischnogaster. The sole species included in Holischnogaster, gracilipes, is evidently the sister-group to Parisch- nogaster, but possesses no described autapomorphies. SUBFAMILY StENOGASTRINAE BEQUAERT, 1918 (ISCHNOGASTERINAE ASHMEAD, 1902) Genus Anischnogaster van der Vecht Anischnogaster van der Vecht, 1972, Zool. Meded. 47:240, 241, genus. Type species: Ischnogaster iridipennis Smith, 1859, by original designation. dubia van der Vecht Anischnogaster dubia van der Vecht, 1972, Zool. Meded. 47:242 (key), 247, figs. 24-36, female, male — “N.W. New Guinea . . . Araucaria Camp, 800 m” (ho- lotype female Leiden); also from other localities in Irian Jay a. Distribution: Indonesia: Irian Jaya. 1996 CHECKLIST OF STENOGASTRINAE 23 iridipennis (Smith) Ischnogaster iridipennis Smith, 1859, J. Proc. Linn. Soc., Zool. 3:166, male — “Am” (lectotype Oxford). — ^Meade-Waldo, 1914, Ann. Mag. Nat. Hist (8)14:405 (syn.: Ischnogaster malayaensis Cameron). — van der Vecht, 1972, Zool. Meded. 47:242 (designation of lectotype). Ischnogaster malayaensis Cameron, 1906, Tijdschr. Ent. 49:231, female — “Etna Bay” (Amsterdam). — Meade- Waldo, 1914, Ann. Mag. Nat. Hist. (8)14:405 (syn. of Isch- nogaster iridipennis Smith). Ischnogaster coriaceus du Buysson, 1907, Notes Leyden Mus. 29:79, male — “Nou- velle Guinee” (Leiden). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):255 (in subgenus Parischnogaster). — van der Vecht, 1972, Zool. Meded. 47:243 (syn. of Anischnogaster iridipennis (Smith)). Stenogaster iridipennis, von Schulthess, 1927, Suppl. Ent. 16:86, 87 (? or Parischno- gaster). Parischnogaster ? coriaceus', von Schulthess, 1927, Suppl. Ent. 16:87. Parischnogaster ? malayaensis', von Schulthess, 1927, Suppl. Ent. 16:87. Anischnogaster iridipennis', van der Vecht, 1972, Zool. Meded. 47:241, 242, figs. 1-13, 57, 58 (syns.: Ischnogaster malayaensis Cameron, Ischnogaster coriaceus du Buysson). Distribution: Indonesia: Am, Misool, Waigeo, Irian Jaya; Papua New Guinea. laticeps van der Vecht Anischnogaster laticeps van der Vecht, 1972, Zool. Meded. 47:242 (key), 249, figs. 37-46, 59, female, male — “North West New Guinea, Araucaria Camp, 800 m” (holotype female Leiden); also from other locafities in New Guinea. Distribution: Indonesia: Irian Jaya; Papua New Guinea. loriai loriai (du Buysson) Ischnogaster loriai du Buysson, 1909, Ann. Mus. Civ. Stor. Nat. Genova 44:313, male — “Nouvelle Guinee S. E. Moroka, 1,300 m” (Genova). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):254 (in subgenus Ischno- gaster) ■ Parischnogaster ? loriai', von Schulthess, 1927, Suppl. Ent. 16:87. Anischnogaster loriai loriai', van der Vecht, 1972, Zool. Meded. 47:242, 251, 253, figs. 47-56, pi. 1. Distribution: Indonesia: Irian Jaya; Papua New Guinea. loriai maculata van der Vecht Anischnogaster loriai maculata van der Vecht, 1972, Zool. Meded. 47:253, female, male — “N. E. New Guinea . . . Mt. Kaindi, Nami Ck., 1700 m” (holotype male Honolulu); also from other localities in Irian Jaya. Distribution: Indonesia: Irian Jaya. spilaspis (Cameron) Ischnogaster spilaspis Cameron, 1913, Bijdr. Dierk. 19:78, male — “Waigeu” (Amsterdam). 24 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Anischnogaster spilaspis; van der Vecht, 1972, Zool. Meded. 47:242 (key), 246, figs. 14-23 (“The original description ... no doubt . . . based on a male” “a female from ‘Waigeoe, 31 Dec.’, labelled ''‘‘Ischnogaster spilaspis Cam. type by Cameron is in Mus. Amsterdam”, but “It appears uncertain whether the true type [male] is still in existence”). Distribution: Indonesia: Waigeo, Irian Jaya. Genus Eustenogaster van der Vecht Eustenogaster van der Vecht, 1969, in Yoshikawa et ah. Nature and Life in S. E. Asia 6:165, genus. Type species: Ischnogaster micans de Saussure, 1852, by original designation. Paravespa Bell, 1936, J. Bombay Nat. Hist. Soc. 38: Fig. 1, 803, 806, genus (2 species). Type species: Paravespa eva Bell, 1936 [= Stenogaster eximioides Dover and Rao, 1922], by subsequent designation of van der Vecht and Carpenter, 1990, Zool. Verh., Leiden 260:44. Junior homonym of Paravespa Radosz- kowski, 1886 (Vespidae: Eumeninae). agilis (Smith) Ischnogaster agilis Smith, 1860, J. Proc. Linn. Soc., Zool. 5, SuppL: 89, male — “Makassar” (Oxford). Stenogaster agilis; von Schulthess, 1927, Suppl. Ent. 16:85. Eustenogaster agilis; Carpenter, 1988, J. New York Ent. Soc. 96:174. Distribution: Indonesia: Sulawesi. calyptodoma (Sakagami and Yoshikawa) Stenogaster calyptodoma Sakagami and Yoshikawa, 1968, Annot. Zool. Japon. 41(2):77, figs. 1-2, 4, 6, female — “Kampong Astana near Kuching, Sarawak, Malaysian Borneo” (Hokkaido University, Sapporo). Eustenogaster calyptodoma; Yoshikawa et al, 1969, in Nature and Life in S E. Asia 6:157. Distribution: Malaysia: Peninsular Malaysia, Sarawak; Indonesia: Sumatra. eximia eximia (Bingham) Ischnogaster eximius Bingham, 1890, J. Bombay Nat. Hist. Soc. 5:244, male — “Ceylon” (London). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):256 (in subgenus Ischnogaster). Ischnogaster eximia; von Dalla Torre, 1904, Genera Insect. 19:83 (cat.). Stenogaster eximia; Dover and Rao, 1922, J. Asiat. Soc. Bengal (N. S.) 18: 241. — Dover, 1925 (1924), J. Asiat. Soc. Bengal (N. S.) 20:301 (syn.: Steno- gaster eximioides Dover and Rao); 1931, J. Fed. Malay St. Mus. 16:256 (syn.: Ischnogaster omatifrons Cameron). Stenogaster eximius; von Schulthess, 1927, Suppl. Ent. 16:85. Eustenogaster eximia; Iwata, 1976, Evol. Instinct: 275 (ethology). — Krombein, 1976, Loria 13(6):303. 1996 CHECKLIST OF STENOGASTRINAE 25 Eustenogaster eximia eximia\ Das and Gupta, 1984 (1983), Or. Ins. 17:401 (cat.). Distribution: Sri Lanka; Thailand; Malaysia: Peninsular Malaysia. eximia eximioides (Dover and Rao) Stenogaster eximioides Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 18:242, male — “Nadgani, Nilgiri Hills. 2,500 ft.” (lectotype Calcutta). — Dover, 1925 (1924), J. Asiat. Soc. Bengal (N. S.) 20:301 (syn. of Stenogaster eximia (Bingham)). — Das and Gupta, 1989, Or. Ins. Monogr. 11:28 (designation of lectotype). Paravespa eva Bell, 1936, J. Bombay Nat. Hist. Soc. 38:803, fig. 1, female, male — “N. Kanara District, Bombay” (London). — Das and Gupta, 1984 (1983), Or. Ins. 17:402 (syn. of Eustenogaster eximia eximioides). Eustenogaster eximioides, Iwata, 1976, Evol. Instinct: 275 (ethology). Eustenogaster eximia eximioides. Das and Gupta, 1984 (1983), Or. Ins. 17:402 (syn.: Paravespa eva Bell; cat.). 'dschnogaster eximia eximioides""', van der Vecht and Carpenter, 1990, Zool. Verh., Leiden 260:44 (designation as type species of Paravespa Bell). Distribution: India: Kerala, Tamil Nadu. fraterna (Bingham) Ischnogaster fratema Bingham, 1897, Fauna Brit. India, Hym. 1:377 (key), 378, female, male — “Tenasserim” (London). Stenogaster fraterna', Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 18:240.— Dover, 1925 (1924), J. Asiat. Soc. Bengal (N. S.) 20:301 (syns.: Isch- nogaster scitula Bingham, Stenogaster scitula var. assamensis Dover and Rao). Eustenogaster fraterna', Iwata, 1976, Evol. Instinct: 275 (ethology). Distribution: India: Assam, Sikkim; Myanmar; Malaysia: Peninsular Malaysia. fulvipennis (Cameron) Ischnogaster fulvipennis Cameron, 1902, J. Straits Branch R. Asiat. Soc. 37:106, female, male — “Mt. Penrissen, Sarawak” (London). — Dover, 1929, Bull. Raf- fles Mus. 2:45 (syn. of Stenogaster micans (de Saussure)). ? Ischnogaster fuscipes Cameron, 1903, J. Straits Branch R. Asiat. Soc. 39:171, sex not stated — “Klackang” (London). Stenogaster fulvipennis', von Schulthess, 1927, Suppl. Ent. 16:86. Eustenogaster fulvipennis'. Carpenter, 1988, J. New York Ent. Soc. 96:175. Distribution: Malaysia: Sarawak. hauxwellii (Bingham) Ischnogaster rufo-maculata Bingham, 1894, J. Bombay Nat. Hist. Soc. 8:385, female — “Tenasserim (Salween Valley, Tavoy)” (London); 1897, Fauna Brit. India, Hym. 1:377, 379 (rufomaculata). — Das and Gupta, 1984 (1983), Or. Ins. 17:402 (syn. of Eustenogaster hauxwellii (Bingham), acting as first reviser). Ischnogaster hauxwellii Bingham, 1894, J. Bombay Nat. Hist. Soc. 8(3):386, pi. 1 fig. 4, female, male — “Tenasserim (Ataran and Ye Valleys)” (London). 26 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Ischnogaster hauxwelli [!]; von Dalla Torre, 1904, Gen. Ins. 19:84 (syn. of Ischno- gaster micans de Saussure). ? Stenogaster bicarinata Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 18:242, male — “Ten miles south of Kuching, Sarawak, Borneo” (Calcutta). Stenogaster rufomaculata; Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 18:241. Stenogaster micans var. hauxwelli [!]; von Schulthess, 1927, Suppl. Ent. 16:85. Stenogaster rufomaculatus; von Schulthess, 1927, Suppl. Ent. 16:86 (cat.). Stenogaster nigromaculata [!]; Dover, 1931, J. Fed. Malay. St. Mus. 16:256. Eustenogaster hauxwellii; Iwata, 1976, Evol. Instinct: 274. — Das and Gupta, 1984 (1983), Or. Ins. 17:402 (syn.: Ischnogaster rufomaculata Bingham, acting as first reviser; cat.; distr.). Distribution: India: Sikkim; Myanmar; Thailand; Malaysia: Sarawak; Singapore; Indonesia: Java, Sumatra*, Bangka*, Kalimantan*; Philippines: Mindanao. luzonensis (Rohwer) Stenogaster micans var. luzonensis Rohwer, 1919, Bull. Hawaii. Sugar Pltrs. As- soc. Exp. Stat. (Ent. Ser.) 14(l):3a, 16, male, female — “Mt. Makiling, Luzon, P. I.” (holotype female Washington). Stenogaster micans var. luzonicus [!]; von Schulthess, 1927, Suppl. Ent. 16:85. Eustenogaster luzonensis^ Carpenter, 1988, J. New York Ent. Soc. 96:175. Distribution: Philippines: Luzon. micans (de Saussure) Ischnogaster micans de Saussure, 1852, Ann. Soc. Ent. France (2)10:24, fe- male— “Java” (Leiden). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):254 (in subgenus Ischnogaster). ? Ischnogaster ornatifrons Cameron, 1902, J. Straits Branch R. Asiat. Soc. 37: 105, female — “Santubong, Kuching, Sarawak” (London). — Meade-Waldo, 1914, Ann. Mag. Nat. Hist. (8)14:463 (in subgenus Parischnogaster). Stenogaster micans; von Schulthess, 1927, Suppl. Ent. 16:81, 85. — Dover, 1929, Bull. Raffles Mus. 2:45 (syn.: Ischnogaster fulvipennis Cameron). Stenogaster ornatifrons; von Schulthess, 1927, Suppl. Ent. 16:86. Eustenogaster micans; van der Vecht, 1969, in Yoshikawa et ah. Nature and Life in S. E. Asia 6:165. Holischnogaster micans; Lee, 1982, Hornets Agric. Regions China: 133 [probable error or misidentification]. Distribution: Myanmar; Thailand; Malaysia: Peninsular Malaysia*, Sarawak; Sin- gapore; Indonesia: Java, Sumatra*, Bangka*. palavanica Reyes Eustenogaster palavanica Reyes, 1988, Philipp. Ent. 7(4):387, 392 (key), figs. 25- 29, female, male — “Irawan River, 2000 ft, Mt. Beaufort, Palawan” (holotype male Honolulu); also from other localities in Palawan. Distribution: Philippines: Palawan. 1996 CHECKLIST OF STENOGASTRINAE 27 scitula (Bingham) Ischnogaster scitula Bingham, 1897, Fauna Brit. India, Hym. 1:377 (key), 379, fe- male— “Rangit Valley, Sikhim; Margherita, Assam” (London). — Dover, 1925 (1924), J. Asiat. Soc. Bengal (N. S.) 20:301 (syn. of Stenogaster fratema (Bing- ham)). Stenogaster scitula; Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 18:240. Stenogaster scitula var. assamensis Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 18:240, male — “Margherita, Assam” (Calcutta). — Dover, 1925 (1924), J. Asiat. Soc. Bengal (N. S.) 20:301 (syn. of Stenogaster scitula (Bing- ham)). Stenogaster seitula [!]; Lee, 1982, Hornets Agric. Regions China: 130. Eustenogaster scitula; Das and Gupta, 1984 (1983), Or. Ins. 17:402 (syn.: Sten- ogaster scitula var. assamensis Dover and Rao). Distribution: India: Assam, Meghalaya, Sikkim; Myanmar; Malaysia. Genus Liostenogaster van der Vecht Liostenogaster van der Vecht, 1969, in Yoshikawa et al, Nature and Life in S. E. Asia 6: 165, genus. Type species: Ischnogaster nitidipennis de Saussure, 1853, by original designation. flaviplagiata (Cameron) Ischnogaster flaviplagiatus Cameron, 1902, J. Straits Branch R. Asiat. Soc. 37: 104, female — “Kuching, Sarawak” (London). Parischnogaster flavoplagiatus [!]; von Schulthess, 1927, Suppl. Ent. 16:87. Stenogaster flavoplagiata [!]; Dover, 1931, J. Fed. Malay. St. Mus. 16:256 (syn. of Stenogaster nitidipennis (de Saussure)). Liostenogaster flaviplagiata; Turillazzi and Carfi, 1996, Trop. Zool. 9(1): 19. Distribution: Thailand; Malaysia: Sarawak; Indonesia: Sumatra. flavolineata (Cameron) Ischnogaster flavolineata Cameron, 1902, J. Straits Branch R. Asiat. Soc. 37:108, female — “Lingga, Sarawak” (London). — Meade-Waldo, 1914, Ann. Mag. Nat. Hist. (8)14:463 (in subgenus Parischnogaster). Ischnogaster butteli von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):255, 257, Figs. F, J-L, male, female (in subgenus Parischnogas- ter)— “Malacca, Taiping Hills, . . . Maxwell’s Hill, Taiping” (Berlin). — Meade- Waldo, 1914, Ann. Mag. Nat. Hist. (8)14:463 (in subgenus Parischnogaster; “This species is distinct from 7. flavolineata Cam.”). — Dover, 1931, J. Fed. Malay. St. Mus. 16:256 (syn. of Stenogaster flavolineata (Cameron)). Parischnogaster butteli; von Schulthess, 1927, Suppl. Ent. 16:86. Parischnogaster flavolineatus; von Schulthess, 1927, Suppl. Ent. 16:87 (distr.). Stenogaster flavolineata; Dover, 1929, Bull. Raffles Mus. 2:45. Liostenogaster flavolineata; Carpenter, 1988, J. New York Ent. Soc. 96:174. Distribution: Malaysia: Peninsular Malaysia, Sarawak; Singapore*; Indonesia: Su- matra. 28 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) lunata Carpenter Liostenogaster lunata Carpenter, 1982, Syst. Ent. 7:34. Nomen nudum. nitidipennis (de Saussure) Ischnogaster nitidipennis de Saussure, 1853, Et. Earn. Vesp. 2:10, male — “Selon r etiquette, Cayenne!?? mais il est evident a mes yeux que cet insecte ne pent venir que de Java, de la Nouvelle-Guinee, ou de la Polynesie” (Paris). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):255 (in subgenus Parisch- nogaster). Parischnogaster nitidipennis', von Schulthess, 1927, Suppl. Ent. 16:88. Stenogaster nitidipennis', Dover, 1929, Bull. Raffles Mus. 2:46; 1931, J. Fed. Malay. St. Mus. 16:256 (syn.: Stenogaster flavoplagiata [!] (Cameron)). Liostenogaster nitidipennis', Iwata, 1976, Evol. Instinct: 275 (ethology). Distribution: Myanmar; Thailand; Malaysia: Peninsular Malaysia, Sabah*; Singapore; Indonesia: Java, Kalimantan, Sumatra*; Philippines: Luzon, Palawan. pardii Turillazzi and Carfi Liostenogaster pardii Turillazzi and Carfi, 1996, Trop. Zool. 9(1): 19, 20, figs. 1-22, male, female, nest — "'Malaysian Peninsula. Gentrng Sempah, Genting Tea Estate (elev. 610 m), Pahang State” (holotype male Firenze); also from Selangor and Sarawak. Distribution: Malaysia: Peninsular Malaysia, Sarawak. picta (Smith) Ischnogaster pictus Smith, 1860, J. Proc. Linn. Soc., Zool. 5, Suppl.: 89, female — “Makassar” (Oxford). Ischnogaster picta', von Dalla Torre, 1894, Cat. Hym. 9:113 (cat.). Stenogaster pictus', von Schulthess, 1927, Suppl. Ent. 16:86. Liostenogaster picta', Turillazzi and Carfi, 1996, Trop. Zool. 9(1): 19. Distribution: Indonesia: Sulawesi. varipicta (Rohwer) Stenogaster varipictus Rohwer, 1919, Bull. Hawaii. Sugar Pltrs. Assoc. Exp. Stat. (Ent. Ser.) 14(l):3a, 15, female — “Los Banos, Luzon, P. I.” (Washington). Parischnogaster ? varipictus', von Schulthess, 1927, Suppl. Ent. 16:88. Liostenogaster varipicta'. Carpenter, 1981 (1982), Syst. Ent. 7:34. Liostenogaster variapicta [!]; Reyes, 1988, Phihpp. Ent. 7(4): 396. Distribution: Thailand*; Malaysia: Peninsular Malaysia, Sabah*; Singapore; Indone- sia: Sumatra*, Kalimantan; Phihppines: Palawan, Visayas, Mindanao, Luzon. vechti Turillazzi Liostenogaster arcuata Hansell, 1987, Oxford Surv. Evol. Biol. 4:167. Nomen nudum. 1996 CHECKLIST OF STENOGASTRINAE 29 Liostenogaster vechti Turillazzi, 1988, Tropic. Zool. 1:193, 194, figs. 1-20, female, male — “Bukit Fraser (Fraser Hill), Jeriau Waterfall (910 m a.s.L), Pahang State” (holotype male Firenze); also from other locahties in Malaysia. Distribution: Malaysia: Peninsular Malaysia; Indonesia: Sumatra. Genus Metischnogaster van der Vecht Metischnogaster van der Vecht, 1977, Tijdschr. Ent. 120:59, 60, genus. Type species: Ischnogaster cilipennis Smith, 1857, by original designation. cilipennis (Smith) Ischnogaster cilipennis Smith, 1857, Cat. Hym. Brit. Mus., Hym. 5:92, male — “Ma- lacca (Mount Ophir)” (lectotype London). — du Buysson, 1909, Ann. Mus. Civ. Stor. Nat. Genova 44:312 (syn.: Ischnogaster drewseni (de Saussure)). — von Schul- thess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):255 (in subgenus Parischnogaster). — van der Vecht, 1977, Tijdschr. Ent. 120:64 (designation of lec- totype). Parischnogaster cilipennis-, von Schulthess, 1927, Suppl. Ent. 16:87 (syn.: Ischno- gaster drewseni de Saussure). Stenogaster cilipennis-, Dover, 1931, J. Fed. Malay St. Mus. 16:256 (distr.). Metischnogaster cilipennis-, van der Vecht, 1977, Tijdschr. Ent. 120:63, 64. Distribution: Malaysia: Peninsular Malaysia, Sarawak; Indonesia: Sumatra, Kaliman- tan. drewseni (de Saussure) Ischnogaster drewseni de Saussure, 1857, Ann. Soc. Ent. France (3)5:315, male — “Poulo-Pinang, ile de TArchipel Indien” (Geneve). — du Buysson, 1909, Ann. Mus. Civ. Stor. Nat. Genova 44:312 (syn. of Ischnogaster cilipennis Smith). — von Schul- thess, 1927, Suppl. Ent. 16:87 (syn. of Parischnogaster cilipennis (Smith)). Ischnogaster drewsenii [!]; von Dalla Torre, 1894, Cat. Hym. 9:113 (cat.). Stenogaster drewseni-, Pagden, 1962, Malay. Nat. J. 16:95 (ethology). Metischnogaster drewseni-, van der Vecht, 1977, Tijdschr. Ent. 120:64 (key), 69. Distribution: Malaysia: Peninsular Malaysia, Sarawak; Singapore; Indonesia: Sumatra, Behtung, Java, Kalimantan; Phihppines: Palawan. Genus Parischnogaster von Schulthess Parischnogaster von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37: 255, subgenus of Ischnogaster Guerin. Type species: Ischnogaster mellyi de Saussure, 1852, by original designation. Holischnogaster van der Vecht, 1977, Tijdschr. Ent. 120:60, 70, genus. Type species: Holischnogaster gracilipes van der Vecht, 1977, by original desig- nation and monotypy. alternata Sakagami ? Stenogaster sarawakensis Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 18:240, female — “10 miles south of Kuching, Sarawak, Borneo” (Calcutta). 30 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Parischnogaster altemata Sakagami, 1969, in Yoshikawa et ah, Nature and Life in S. E. Asia 6:155, female, male — “Fraser’s Hill, Malaya” (holotype male Hokkaido University, Sapporo); also from another locality in Peninsular Malay- sia. Parischnogaster alternate [!]; Kojima, 1990, Japan. J. Ent. 58(3):517. Distribution: Thailand*; Malaysia: Peninsular Malaysia, Sabah, Sarawak; Singa- pore*; Indonesia*: Sumatra. aurifrons (Smith), NEW COMBINATION Ischnogaster aurifrons Smith, 1862, J. Proc. Linn. Soc., Zool. 6:58, female — “Celebes (Tondano)” (Oxford). Stenogaster aurifrons; von Schulthess, 1927, Suppl. Ent. 16:85, 86 (? or Paris- chnogaster). Distribution: Indonesia: Sulawesi. depressigaster (Rohwer) Stenogaster depressigaster Rohwer, 1919, Bull. Hawaii. Sugar Pltrs. Assoc. Exp. Stat. (Ent. Ser.) 14(l):3a, 16, male — “Los Banos, Luzon, P. I.” (Washington). Parischnogaster ? dpressigaster [!]; von Schulthess, 1927, Suppl. Ent. 16:87. Parischnogaster depressigaster; Iwata, 1967, Nature Life S. E. Asia 5:260 (ethol- ogy). Distribution: Philippines: Luzon. gracilipes (van der Vecht) Holischnogaster gracilipes van der Vecht, 1977, Tijdschr. Ent. 120:73, figs. 38- 49, female, male — “Borneo: Sabah . . . Mt. Kinabalu, 5,000 ft” (Venezia); also from other localities in Borneo and Sumatra. Parischnogaster gracilipes; Carpenter, 1988, J. New York Ent. Soc. 96:175. Distribution: Malaysia: Sabah, Sarawak; Indonesia: Sumatra. jacobsoni (du Buysson) Ischnogaster jacobsoni du Buysson, 1913, Bull. Mus. Hist. Nat. Paris 7:436, male — “Java, Noesa Kambangan” (Paris). Parischnogaster jacobsoni; von Schulthess, 1927, Suppl. Ent. 16:81, 87. Distribution: Malaysia: Peninsular Malaysia, Sabah; Indonesia: Sumatra, Java, Bangka*, Kalimantan*. mellyi (de Saussure) Ischnogaster mellyi de Saussure, 1852, Ann. Soc. Ent. France (2)10:25, pi. 2, fig. 1, male — “Java” (male Geneve). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):256 (in subgenus Parischnogaster). Ischnogaster nigrifrons Smith, 1857, J. Proc. Linn. Soc., Zool.: 113, female — “Borneo (Sarawak)” (Oxford). — Das and Gupta, 1984 (1983), Or. Ins. 17:401 (syn. of Parischnogaster mellyi (de Saussure)). Stenogaster nigrifrons; Dover and Rao, 1922, J. Proc. Asiat. Soc. Bengal (N. S.) 1996 CHECKLIST OF STENOGASTRINAE 31 18:241.— Dover, 1929, Bull. Raffles Mus. 2:45; 1931, J. Fed. Malay St. Mus. 16:255 (syns.: Ischnogaster nigricans Cameron, Ischnogaster serrei du Buys- son). Parischnogaster mellyi; von Schulthess, 1927, Suppl. Ent. 16:82, 87. — Das and Gupta, 1984 (1983), Or. Ins. 17:401 (syn.: Ischnogaster nigrifrons Smith). Parischnogaster nigrifrons’, von Schulthess, 1927, Suppl. Ent. 16:83, 87. Stenogaster mellyi’, Pagden, 1962, Malay. Nat. J. 16:95. Distribution: India: Assam, Meghalaya, Sikkim; Myanmar; Vietnam; Thailand; Malaysia: Peninsular Malaysia, Sabah; Singapore; Indonesia: Sumatra, Java, Ka- limantan, Bangka*; Philippines: Mindanao. nigricans nigricans (Cameron) Ischnogaster nigricans Cameron, 1902, J. Straits Branch R. Asiat. Soc. 37:105, female — “Kuching, Sarawak” (London). — Meade-Waldo, 1914, Ann. Mag. Nat. Hist. (8)14:463 (in subgenus Parischnogaster). — Dover, 1931, J. Fed. Malay St. Mus. 16:255 (syn. of Stenogaster nigrifrons (Smith)). Parischnogaster nigricans’, von Schulthess, 1927, Suppl. Ent. 16:87. Distribution: Malaysia: Peninsular Malaysia, Sarawak; Singapore*; Indonesia: Su- matra*, Bangka*, Krakatau*, Java*, Kangean*; Philippines: Luzon. nigricans serrei (du Buysson) Ischnogaster serrei du Buysson, 1905, Bull. Soc. Ent. France: 281, female, male — “Batavia” (Paris). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):255 (in subgenus Parischnogaster). — Dover, 1931, J. Fed. Malay St. Mus. 16:255 (syn. of Stenogaster nigrifrons (Smith)). Parischnogaster serrei’, von Schulthess, 1927, Suppl. Ent. 16:88 (cat.). Stenogaster serripes [!]; von Schulthess, 1935, Rev. Suisse Zool. 42:299. Distribution: Indonesia: Java striatula (du Buysson) Ischnogaster striatulus du Buysson, 1905, Bull. Soc. Ent. France: 282, female — “Sumatra” (Paris). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):255 (in subgenus Parischnogaster). ? Ischnogaster clypealis Cameron, 1906, J. Straits Branch R. Asiat. Soc. 46:122, female — “Bidi . . . Kuching” (London). Ischnogaster foveatus du Buysson, 1907, Notes Leyden Mus. 29:80, male — “Sal- awatti” (Leiden). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):255 (in subgenus Parischnogaster). ? Ischnogaster levifoveatus Meade-Waldo, 1914, Ann. Mag. Nat. Hist. (8)14:462, male (in subgenus Parischnogaster) — “BORNEO: Sarawak, Mt. Matang” (Lon- don). ? Parischnogaster costulatus von Schulthess, 1927, Suppl. Ent. 16:82, female — “Sumatra, Lubuk Sulasih, 1,000 m” (Wien). Parischnogaster striatulus’, von Schulthess, 1927, Suppl. Ent. 16:82 {syn.’. foveatus du Buysson). 32 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Parischnogaster striatulo [!]; von Schulthess, 1927, Suppl. Ent. 16:82. Parischnogaster striolatus [!]; von Schulthess, 1927, Suppl. Ent. 16:83. Parischnogaster foveatus; von Schulthess, 1927, Suppl. Ent. 16:87, 88 (syn.: Par- ischnogaster striatulus (du Buysson)). Parischnogaster ? levifoveatus; von Schulthess, 1927, Suppl. Ent. 16:87. Distribution: Thailand; Malaysia: Peninsular Malaysia*, Sarawak; Indonesia: Su- matra, Kalimantan, Salawati, Java*. timida (Williams) ? Ischnogaster strandi du Buysson, 1910, Bull. Soc. Ent. France: 39, male, fe- male— “Philippines: Samar” (Berlin). Parischnogaster strandi; von Schulthess, 1927, Suppl. Ent. 16:88. Stenogaster timidus Williams, 1928, Bull. Hawaii. Sugar Pltrs. Assoc. Exp. Stat. (Ent. Ser.) 19:170, pi. XXXIII Fig. 228, female— “Molawin Falls, Mt. Makiling, Los Banos” (Honolulu). Parischnogaster timida; Carpenter, 1988, J. New York Ent. Soc. 96:175. Distribution: Philippines: Samar, Luzon. unicuspata Reyes Parischnogaster unicuspata Reyes, 1988, Philipp. Ent. 7(4): 387, 400 (key), figs. 1, 69-75, 82, 85, 91-92, 109, female, male — “Palawan National Agricultural College, Aborlan, Palawan” (holotype male Los Banos); also from other local- ities in Palawan and Malaysia. Distribution: Thailand*; Malaysia: Peninsular Malaysia, Sarawak; Singapore*; In- donesia*: Sumatra, Bangka; Philippines: Palawan. Genus Stenogaster Guerin-Meneville Stenogaster Guerin-Meneville, 1831, in Duperrey, Voyage Coquille, ZooL, Ins., At- las: pi. 9 fig. 9, genus. Type species: Stenogaster fulgipennis Guerin-Meneville, 1831, by monotypy. Ischnogaster Guerin-Meneville, 1838, in Duperrey, Voyage Coquille, ZooL, Ins., Texte 2(1):268. Unnecessary replacement name for Stenogaster Guerin-Mene- ville, 1831, thought to be preoccupied by Stenogaster Solier, 1835 (Coleoptera). adusta van der Vecht Stenogaster adusta van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 287 (key), 291, figs. 21-33, female, male — “Misool . . . Fakal” (holotype female Leiden). Distribution: Indonesia: Misool, Irian Jaya. canaliculata (Cameron) Ischnogaster canaliculatus Cameron, 1911, Nova Guinea 9(2), ZooL: 190, fe- male— “Bivak Island” (lectotype Amsterdam). — Meade-Waldo, 1914, Ann. Mag. Nat. Hist (8)14:405 (syn. of Ischnogaster fulvipennis [!] Guerin-Mene- ville).— van der Vecht, 1975, J. Aust. Ent. Soc. 14:299, 301 (desgination of lectotype). 1996 CHECKLIST OF STENOGASTRINAE 33 Stenogaster ? canaliculatus; von Schulthess, 1927, Suppl. Ent. 16:85. Stenogaster canaliculata; van der Vecht, 1975, J. Aust. Ent. Soc. 14:298 (key), 299. Distribution: Indonesia: Irian Jaya. concinna van der Vecht Stenogaster concinna van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 285, 286 (key), 295, figs, la-b, 53-80, male, female — “S.W. New Guinea . . . Wataikwa River” (holotype male London); also from other localities in New Guinea. Distribution: Indonesia: Irian Jaya; Papua New Guinea. decorata van der Vecht Stenogaster decorata van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 287 (key), 293, figs. 34-51, female, male — “Misool . . . Fakal” (holotype female Leiden). Distribution: Indonesia: Misool, Yapen. flavifrons van der Vecht Stenogaster flavifrons van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 298 (key), 303, figs. 94-103, female, male — “N.W. New Guinea . . . Japen, Mt. Baduri, 300 m” (London). Distribution: Indonesia: Yapen. fulgipennis (Guerin-Meneville) Stenogaster fulgipennis Guerin-Meneville, 1831, in Duperrey, Voyage Coquille, Zook, Ins., Atlas: pi. 9 fig. 9, sex and locality not stated (holotype male from “Dory, N. Guinea,” Genova). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37(3):254 (in subgenus Ischnogaster). — Dover and Rao, 1922, J. Asiat. Soc. Beng. (N. S.) 18:236, footnote (correct generic name). — Bequaert, 1926, Ent. Mitt. 15:192 (generic name). — Guiglia, 1948, Ann. Mus. Civ. Stor. Nat. Giacomo Doria 63:178 (location of type). Ischnogaster fulgipennis', Guerin-Meneville, 1938, in Duperrey, Voyage Coquille, Zool., Ins., Texte 2(I):269, “female” from “Dory, dans la Nouvelle-Guinee” [the description is evidently based on a male]. — de Saussure, 1852, Ann. Soc. Ent. France (2)10:23 “fern, ou ouvriere” [redescription of Guerin-Meneville’s type]; 1853, Et. Earn. Vesp. 2:7 (“female”). — von Schulthess, 1914, Zool. Jahrb. (Abt. Syst. Geogr. Biol. Tiere) 37:254 (type of subgenus Ischnogaster). Stenogaster fulgidipennis', von Dalla Torre, 1894, Cat. Hym. 9:113, invalid emen- dation. Ischnogaster fulvipennis [!]; Meade-Waldo, 1914, Ann. Mag. Nat. Hist (8)14:405 (syn.: Ischnogaster canaliculatus Cameron). Distribution: Indonesia: Irian Jaya. glabra van der Vecht Stenogaster glabra van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 299 (key), 303, figs. 104-110, male — “N.W. Guinea . . . Cyclops Mts., 1,100 m” (London). 34 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Distribution: Indonesia: Irian Jaya. licina van der Vecht Stenogaster licina van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 287 (key), 295, fig. 52, female — “N.W. Guinea . . . Road Ajamaru-Teminabuan, km 1-8” (Lei- den). Distribution: Indonesia: Irian Jaya. macilenta van der Vecht Stenogaster macilenta van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 298 (key), 306, figs. Ic-e, 117-131, female, male — “N.W. New Guinea . . . Lower Mist Camp, 1550 m” (holotype female Leiden); also from other localities in Irian Jaya. Distribution: Indonesia: Irian Jaya. pilosa van der Vecht Stenogaster pilosa van der Vecht, 1975, J. Aust. Ent. Soc. 14:283, 299 (key), 305, figs. 111-116, male — “N.W. New Guinea . . . Cyclops Mts, Mt. Lina, 1,100- 1,400 m” (London). Distribution: Indonesia: Irian Jaya. unicolor (Smith) Ischnogaster unicolor Smith, 1864, J. Proc. Linn. Soc., Zool. 7:41, male — “Mysol; Waigiou” (lectotype Oxford). Stenogaster unicolor, von Schulthess, 1927, Suppl. Ent. 16:86. Distribution: Indonesia: Waigeo, Misool. ACKNOWLEDGMENTS We thank C. K. Starr and S. Turillazzi for suggestions, and C. O’Toole, C. van Achterberg, D. Notton, S. Lewis, and J. Casevitz-Weulersse for assistance in examining types. The Neth- erlands Organization for Scientific Research and the Japan Society for the Promotion of Science supported JK’s stay in the Netherlands during the course of this work. LITERATURE CITED Carpenter, J. M. 1981 (1982). The phylogenetic relationships and natural classification of the Vespoidea (Hymenoptera). Syst. Entomol. 7:11-38. Carpenter, J. M. 1988. The phylogenetic system of the Stenogastrinae (Hymenoptera: Vespi- dae). J. New York Entomol. Soc. 96:140-175. Das, B. P. and V. K. Gupta. 1984 (1983). A catalogue of the families Stenogastridae and Vespidae from the Indian Subregion (Hymenoptera: Vespoidea). Orient. Ins. 17:395- 464. Das, B. P. and V. K. Gupta. 1989. The social wasps of India and the adjacent countries (Hymenoptera: Vespidae). Orient. Ins. Monogr. 11:1-292. Hansell, M. H. 1987. Nest building as a facilitating and limiting factor in the evolution of 1996 CHECKLIST OF STENOGASTRINAE 35 eusociality in the Hymenoptera. In: P. H. Harvey and L. Partridge (eds.), Oxford Surveys in Evolutionary Biology 4:155-181. Oxford Univ. Press, Oxford. Iwata, K. 1971. Honno no Shinka: Hachi no Hikaku Shuseigakuteki Kenkyu. Mano Shoten, Kanagawa Prefecture. Iwata, K. 1976. Evolution of Instinct: Comparative Ethology of Hymenoptera. Amerind Pub- lishing Co., New Delhi. Krombein, K. V. 1976. Eustenogaster a primitive social Sinhalese wasp. Loris 13:303-306. Reyes, S. G. 1988. A review of Philippine Stenogastrinae (Hymenoptera: Vespidae). Philipp. Entomol. 7(4):387-434. Schulthess, A. von. 1927. Fauna sumatrensis. (Beitrag Nr. 52). Vespidae (Hym). Suppl. En- tomol. 16:81-92. Turillazzi, S. 1988. Adults and nest of Liostenogaster vechti n. sp. (Hymenoptera, Stenogas- trinae). Tropic. Zool. 1:193-201. Turillazzi, S. 1991. The Stenogastrinae. In: K. G. Ross, and R. W. Matthews (eds.). The Social Biology of Wasps: 74-98. Cornell University Press, Ithaca. Turillazzi, S. and S. Carfi. 1996. Adults and nest of Liostenogaster pardii n. sp. (Hymenoptera Stenogastrinae). Trop. Zool. 9(1): 19-30. Vecht, J. van der. 1969. In: Yoshikawa, et al. Nature and Life in Southeast Asia 6:165. Vecht, J. van der. 1972. A review of the new genus Anischnogaster in the Papuan region (Hymenoptera, Vespidae). Zool. Meded. 47:240-256. Vecht, J. van der. 1975. A review of the genus Stenogaster Guerin (Hymenoptera: Vespoidea). J. Aust. Entomol. Soc. 14:283-308. Vecht, J. van der. 1977. Studies of Oriental Stenogastrinae (Hymenoptera: Vespoidea). Tijdschr. Ent. 120:55-75. Yoshikawa, K., R. Ohgushi, and S. E Sakagami. 1969. Preliminary report on entomology of the Osaka City University 5th Scientific Expedition to Southeast Asia 1966. With de- scriptions of two new genera of stenogasterine [sic] wasps by J. van der Vecht. Nature and Life in Southeast Asia 6:153-182. Received 3 June 1996; accepted 23 August 1996. 36 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) INDEX adusta 32 agilis 24 altemata 29 alternate 30 Anischnogaster 22 apressigaster 30 arcuata 28 assamensis 27 aurifrons 30 bicarinata 26 butteli 27 calyptodoma 24 canaliculata 32 canaliculatus 32 cilipennis 29 clypealis 31 concinna 33 coriaceus 23 costulatus 31 decorata 33 depressigaster 30 drewseni 29 drewsenii 29 dubia 22 Eustenogaster 24 eva 25 eximia 24 eximioides 25 eximius 24 flavifrons 33 flaviplagiata 27 flaviplagiatus 27 flavolineata 27 flavolineatus 27 flavoplagiata 27 flavoplagiatus 27 foveatus 31 fraterna 25 fulgidipennis 33 fulgipennis 33 fulvipennis 25, 33 fuscipes 25 glabra 33 gracilipes 30 hauxwelli 26 hauxwellii 25 Holischnogaster 29 iridipennis 23 Ischnogaster 32 jacobsoni 30 laticeps 23 levifoveatus 31 licina 34 Liostenogaster 27 loriai 23 lunata 28 luzonensis 26 luzonicus 26 macilenta 34 maculata 23 malayaensis 23 mellyi 30 Metischnogaster 29 micans 26 nigricans 31 nigrifrons 30 nigromaculata 26 nitidipennis 28 omatifrons 26 palavanica 26 Paravespa 24 pardii 28 Parischnogaster 29 picta 28 pictus 28 pilosa 34 rufo-maculata 25 rufomaculata 25 rufomaculatus 26 sarawakensis 29 scitula 27 seitula 27 serrei 31 serripes 31 spilaspis 23 Stenogaster 32 strandi 32 striatula 31 striatulo 32 striatulus 31 striolatus 32 timida 32 timidus 32 unicolor 34 unicuspata 32 variapicta 28 varipicta 28 varipictus 28 vechti 28 J. New York Entomol. Soc. 104(l-2);37-47, 1996 TYPES OF NEOTROPICAL SPECIES OF SCATOPSIDAE (DIPTERA: PSYCHODOMORPHA). I. EDWARDS’ CHILEAN AND SOUTHERN ARGENTINEAN SPECIES Dalton de Souza Amorim’ and Jean-Paul HaennP Depto. de Biologia, FFCLRP/USP, Av. Bandeirantes 3900, 14040-901 Ribeirao Preto SP, BRAZIL. Abstract. — The types of Edwards’ species of Chilean and southern Argentinean Scatopsidae are redescribed and illustrated: Ectaetia cornuta Edwards, Scatopse setifera Edwards, Scatopse tubifera Edwards, Scatopse constricta Edwards, and Anapausis fuscinervis (Edwards). Notes on each type are added, and the generic position of each species is discussed. Ectaetia cornuta and Anapausis fuscinervis maintain their present generic placements. New combinations are proposed for Pararhexosa tubifera (Edwards), n.comb. and Neorhegmoclemina constricta (Ed- wards), n.comb. The holotype of Scatopse tubifera, described as a male, is actually a female. A new genus — Holoclema, n.gen. — is erected to include Scatopse setifera (Edwards). Holocle- ma is considered to be the sister group of the remaining Rhegmoclematini. The Neotropical species Rhexosa panamensis (Cook) and Rhexosa auricularis (Duda) are transferred to the formerly monotypic genus Abrhexosa Freeman, originally described for a single Oriental spe- cies. Additional distribution data for some species is supplied. S. maritima and S. costaricana are considered incertae sedis in Scatopsinae until their types can be examined. Key words'. Scatopsidae, Neotropical region, types, Holoclema, phylogeny. The Scatopsidae have 42 described species assigned for the Neotropical region. About 120 other, undescribed species are known for the region. Of the described species, three are cosmopolitan, namely Scatopse notata (Linnaeus), Coboldia fuscipes (Meigen), and Holoplagia guamensis (Johannsen). Another four are considered unrecognizable and their types are probably lost (Cook, 1967) — Scatopse fulvitarsis Macquart of Lynch Arribalzaga, Scatopse hyalinata Phillippi, Scatopse pallidipes Lynch Arribalzaga, and Scatopse transatlantica PhiUippi. The remaining 35 Neotropical species of Scatopsidae are placed in the Ectaetiinae, Psectrosciarinae and Scatopsinae. Two belong to Ectaetia, six to Psectrosciara, one to Anapausis, one to Diamphidicus, one to Rhegmoclema, one to Aldrovandiella, three to Rhegmoclemina, six to Neorhegmoclemina, one to Colobostema, one to Holoplagia, and four to Swammerdamella. Four other Neotropical species included by Cook (1967) in Rhexosa do not belong to the genus: Cook (1976) erected Akorhexoza to include the Neotropical species Scatopse dampfi Duda and Akorhexoza cactivora Cook. The struc- ture of the male terminaha indicates that Rhexosa panamensis Cook should be placed in Freeman’s (1985) genus Abrhexosa, also probably the best place for Scatopse auri- cularis Duda. Both these species are herein transferred to this genus. Rhexosa lobata Cook, 1956 is best alotted to a new genus, with a number of other undescribed Neo- ' e-mail dsamorim@usp.br ^ Musee d’Histoire Naturelle, rue des Terreaux 14, CH-2000 Neuchatel. 38 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) tropical species, defined by a considerably modified tergite 9, divided into a pair of longitudinal strips by a mesal incision, and by a mesal perforation on the aedeagus. Cook’s (1967) catalogue of the Neotropical Scatopsidae kept five species in the genus Scatopse: S. notata (Linnaeus), Scatopse fuscipes Meigen, Scatopse costaricana Duda, Scatopse maritima Duda, and Scatopse tubifera Edwards. Scatopse notata does comprise a monophyletic group with S. lapponica, S. alpestris, S. chinensis, and S. globulicaudata, the small taxon to which Scatopse is now restricted. This taxon is typically Holarctic and the presence of the synantropic species S. notata in other regions is most probably due to human transportation. Scatopse fuscipes Meigen was correctly transferred by Cook (1974) to Coboldia, a Swammerdamellini genus. The descriptions of the other three Neotropical species included by Cook (1967) in Scatopse strongly suggest that they do not belong to Scatopse sensu Cook (1974). However, the original descriptions are insufficient to allow a generic replacement and only the re-examination of the types can solve the problem. The types of S. tubifera were re-examined and are redescribed in this paper. Scatopse maritima and S. costaricana are best considered incertae sedis in the Scatopsinae before the types can be re-examined. This first paper in a series redescribing the Neotropical scatopsids deals with Edwards’ (1930) types of Chilean Scatopsidae. The suprageneric, phylogenetic classification of the Scatopsidae of Amorim (1994) is followed. The “groups” artifact is employed to refer to unnamed monophyletic groups in the se- quenced classification of the family (Amorim, 1982a). Many of the phylogenetic infer- ences used here are based on the analysis of the family in Amorim (1982b, MS). Ectaetiinae Ectaetia comuta Edwards Figs. 1-5 Ectaetia comuta Edwards, 1930: 98, pi. X, figs. 11, 12 (wing, male terminalia). Type- locality: CHILE, Llanquihue, Casa Pangue. Types, male lectotype, one male paralec- totype and five female paralectotypes in BMNH; Cook, 1967: 5 (catalogue). The type series of Ectaetia comuta included seven specimens: a male and a female specimen on the same pin and five other specimens on separate pins. The pin with two specimens presents six levels, here described beginning from the top: (1) a strip with the specimens glued to it; (2) “Casa Pangue/4-10.xii.l926”, printed; (3) a plastic strip with the male terminaha dissected in balsam; (4) a rounded red label with “Type” printed; (5) "'Ectaetia! comuta Edw.” handwritten and “E W. Edwards/det. 1930” print- ed; (6) “S. CHILE/Llanquihue Prov./E & M. Edwards/B.M. 1927-63” printed. The only male paratype has labels 2 and 6. The four female paratypes have the label “Castro/ 20-22.xi.l920”, and label 6. We add here some details to the original description, with drawings of the male terminalia (Figs. 1-5). Flagellum with 10 flagellomeres. Meral setae restricted to the dorsal margin of the sclerite. No strong scutellars. Katepistemum apparently bare. Apex of hind tibia slightly flattened, hind femur with a mesal area less well sclerotized. Wing membrane devoid of macrotrichia. The posterior wing veins translucent; C is consider- ably short, with Rg curved. 1996 EDWARDS’ TYPES OF NEOTROPICAL SCATOPSIDAE 39 Figs. 1-5. Ectaetia comuta Edwards. Male holotype. 1. Terminalia, ventral view; 2. Tergite DC, dorsal view; 3. Gonostyle, ventral view; 4. Paramere, ventral view; 5. Aedeagus, ventral view. Psectrosciarinae Anapausis fuscinervis (Edwards) Figs. 6-7 Psectrosciara fuscinervis Edwards, 1930:97, pi. X, Figs. 11, 12 (male terminalia, wing). Type-locality: CHILE, Llanquihue, Casa Pangue; Cook, 1965:16, Figs. 42, 43, 45 (male terminalia, wing); Cook, 1967:5 (catalogue); Amorim, 1982b (mouth parts). A single male specimen is the only type of this species. The pin with the male holotype has seven labels. Beginning at the top: the specimen glued to a plastic strip; the second with the wing glued to a plastic strip; the third is a printed label written “Casa Pangue/4-10.xii.l926”; the fourth has the terminalia dissected in balsam on a plastic strip; the fifth is a red, round label with “Type” printed; the sixth is a label with "'Psectrosciara! fuscinervis Edw.” handwritten and “E W. Edwards/ 1930” printed; the last label has “S. CHILE/Llanquihue Prov./E & M. Edwards/B.M. 1927-63” printed. In some details, neither Edwards’ (1930) nor Cook’s (1965) drawings are sufficient to allow a correct understanding of the homology of the male terminalia sclerites, although they are sufficient to allow the correct identification of the species. The male terminalia are redrawn here in ventral and dorsal aspect (Figs. 6-7). Some descriptive information may be added: the meron is well separated from epistemum IQ, this latter sclerite well 40 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Figs. 6-7. Anapausis fuscinervis (Edwards). Male holotype. 6. Terminalia, ventral view; 7. Terminalia, dorsal view. developed; tarsi brown; antenna with 8 flagellomeres; wing membrane with abundant macrotrichia present on cells posterior to M2, but absent anteriorly to M2. Other locahties are known for this species: one male, “Puerto Puyuhuapi, Sud Chile, 14-11-39, G. H. Schwabe” (Cook, 1965); a female, Chile, El Naranjo, Tilana, Coquim- bo, X.1967, L. E. Pena col. (Natural History Museum, London) (Amorim, 1982b); one male and one female, Chiloe, Isla Chiloe, Ahoni Alto, September 1988, Luis E. Pena col. (Carnegie Museum of Natural History, Pittsburgh); Castro, Isla Chiloe, xii.1926, R & E Shannon col. (National Museum of Natural History, Washington, D.C.). Scatopsinae Rhegmoclematini Neorhegmoclemina constricta (Edwards), new combination Figs. 8-12 Scatopse constricta Edwards, 1930:96, pi. DC, Figs. 5, 7 (male terminaha, wing). Type-locality: Chile, Llanquihue, Peulla. Rhegmoclemina {Neorhegmoclemina) constricta. Cook, 1967:3 (catalogue); Amorim, 1982b (palpus, thorax, wing, male terminalia). Edwards (1930) described N. constricta based on ten specimens from the same lo- cahty. The male holotype pin has seven layers of labels: (1) the highest one is a plastic strip with some broken parts of the body; (2) a label with handwritten “in log with/ termites”; (3) a label “Peulla/12-13.xii.l926”, printed; (4) a plastic strip with the ter- minaha dissected in balsam; (5) a round, red label with “Type” printed; (6) '"Sca- topse! constricta Edw.” handwritten on a label with “E W. Edwards/det. 1930” printed; (7) a label with “S. CHILE/Llanquihue Prov./E & M. Edwards/B.M. 1927-63” printed. The type series also includes one female with labels 1-4 and 7, one male with labels 1-3 and 7, and seven males with labels 1, 3 and 7. Cook (1955) only stated that it was probable that Scatopse constricta Edwards belongs to Rhegmoclemina, without formally including it in the genus. In the catalogue of the 1996 EDWARDS’ TYPES OF NEOTROPICAL SCATOPSIDAE 41 Figs. 8-10. Neorhegmoclemina constricta (Edwards), n.comb. Male holotype. 8. Segment VII, ventral view; 9. Terminalia, dorsal view; 10. Gonostyle, ventral view. family for the Neotropical region (Cook, 1967) the species was formally transferred to R. {Neorhegmoclemina). Amorim (1994) raised Neorhegmoclemina and Austroclemina to genera and the species is here formally transferred to Neorhegmoclemina. Amorim (1982b) studied four additional males belonging to this species, from Chile (Chepu, Chiloe, 42°S, 20.x. 1958, Royal Society Chilean Expedition [Natural History Museum]), and two other undescribed species of Neorhegmoclemina from Chile (one in the Museu de Zoologia, Sao Paulo, the other in the Canada National Collection). The male termin- aha of the holotype is redrawn here (Fig. 8-10) to avoid any doubt on the identification of the species. The terminafia of the female paratype is figured for the first time (Figs. 11-12). Figs. 11-12. Neorhegmoclemina constricta (Edwards), n. comb. Female paratype. 11. Ter- minalia, ventral view; 12. Spermatheca. 42 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Holoclema, new genus Type-species, Scatopse setifera Edwards, 1930. The following combination of features is diagnostic for this genus; antenna with 10 flagellomeres; palpus small; R5 long, parallel but not close to C; Mj complete; CUA2 rather sinuous, with some macrotrichia; membrane posterior to CUA2 with macrotrichia; a fold between the medial fork and CuAj; tergite 1 without reticulation; gonocoxites with macrotrichia, well developed laterally; parameres absent; aedeagus quite wide; grad- ually reducing in width; a pair of spiracles dorsally on the male terminaha; no cerci. Species included: Holoclema setifera (Edwards, 1930). Holoclema setifera (Edwards), new combination Figs. 13-20 Scatopse setifera Edwards, 1930:95, pi. IX, Figs. 3-4 (wing, male terminaha). Type- locahty: ARGENTINA, Rio Negro, Lago Correntoso. Rhegmoclema setifera'. Cook, 1967:3 (catalogue). The holotype of this species has six levels of labels on the pin: (1) the specimen double-mounted on a plastic strip; (2) a label printed “L. Correntoso/18-25.xi.l926”; (3) a plastic strip with the male terminaha dissected in balsam; (4) a round, red label printed “Type”; (5) a label with ''Scatopse/ setifera Edw.” handwritten and “E W. Ed- wards/det. 1930” printed; (6) “ARGENTINA/rerr. Rio Negro/E & M. Edwards/B.M. 1927-63” printed on a label. Edwards (1930:95) refers to 12 paratypes. All they come from the type locahty and show labels 1, 2, 5 and 6. The type series includes 5 males, 5 females, a wing remaining from a lost specimen, plus an empidid specimen, what would be unexepected for Edwards’ type series. Segment 7 of the holotype is drawn here for the first time (Fig. 13) and the male terminaha are redrawn (Figs. 14-18). Terminaha of the dissected female paratype are also illustrated (Figs. 19-20). Discussion: The question of the phylogenetic position of Holoclema setifera is quite difficult. Edwards (1930) rejected most of Enderlein’s (1912) genera and this species was maintained in the genus Scatopse, as most of his Scatopsinae species. In the Neo- tropical catalogue of the Scatopsidae, Cook (1967) placed the species in the genus Rhegmoclema, but the reexamination of the type shows that this is certainly not satis- factory. The inclusion of Holoclema in the Scatopsinae seems unquestionable: an independent ejaculatory apodeme — an outstanding autapomorphic feature of the subfamily — ^is pres- ent (Edwards, 1930, pi. IX, hg. 4). The decision on the tribal placement, however, deserves careful discussion. The presence of 10 flagellomeres and the presence of ma- crotrichia on the posterior wing veins definitely excludes it from the Scatopsini+ (which includes Scatopsini, Colobostematini and Swammerdamellini). Edwards (1930) associ- ated the species with S. nigripennis, now included in Colobostema, possibly based on the following features: “shght and uniform smoky tinge” color of the wing membrane, “posterior veins somewhat darkened”, rather longer pubescence, and eyes in contact above the antennae only for a small distance. In fact, these are similarities shared with some Colobostematini, but Holoclema is plesiomorphic for a number of synapomorphies of the Scatopsini+, of the Colobostematini + and of the Colobostematini themselves. 1996 EDWARDS’ TYPES OE NEOTROPICAL SCATOPSIDAE 43 Figs. 13-20. Holoclema setifera (Edwards), n.comb. Male holotype. 13. Segment VII, ven- tral view; 14. Terminalia, ventral view; 15. Tergites VIII + IX, ventral view; 16. Gonostyle, ventral view; 17. Right gonocoxite, ventral view; 18. Aedeagus, ventral view. Female paratype; 19. Terminalia, ventral view; 20. Terminalia, dorsal view. suggesting that a hypothesis of close relationship with Colobostema is not parsimonious, and these similarities should be considered homoplastic. Other features are relevant in the taxonomic discussion: (1) 10 flagellomeres; (2) flagellum long; (3) body pubescence long; (4) Mj complete; (5) fork of M long; (6) C 44 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) considerably long; (7) R5 separated from C; (8) R5 parallel to C; (9) macrotrichia on CuA2 and on membrane posterior to CUA2; (10) long scutellar setae present; (11) tergite 1 without reticulation; (12) pretergite 2 hardly differentiated. That there is no reticulation on tergite 1 would be sufficient reason to consider not including the species in Rhegmoclema. Also, some Afrotropical and Oriental species of Rhegmoclema have a complete Mj, but no Neotropical species. Moreover, there is scarce- ly similarity in the male terminaha of Holoclema and Rhegmoclema. Its inclusion in Neorhegmoclemina also seems inappropriate. The absence of macro- trichia on the wing membrane is a synapomorphy of this genus and there are abundant macrotrichia on the proximal-posterior cell in H. setifera Edwards. Also, Parascatopsina has a short R5, a feature definitely absent in Edwards’ species (actually, only Diamphi- dicus and Parmaferia in the Rhegmoclematini show such long R5). Other wing features would show some similarity with Rhegmoclemina or Austroclemina, but only on the basis of plesiomorphies. In addition, Rhegmoclemina and Austroclemina possess an apo- morphic, 8-articulated flagellum, not found in Holoclema. A more detailed study of the male terminaha gives some other clues about its position: it is plesiomorphic for the bowl-shaped, fused gonocoxites, apomorphic in all Rheg- moclematini but Diamphidicus. Tergite 7 is rather well developed, plesiomorphic in H. setifera, differently from the condition presented by the Parascatopsina species. Also important is that the gonocoxites in this species have setae laterally. Gonocoxal setae are absent in all Scatospinae but Diamphidicus. The inclusion of Scatopse setifera Ed- wards in Diamphidicus, however, would also be unsuitable. A long vein R5 and a slender connection between the gonocoxites mesally are features shared by both genera, but they are plesiomorphies; another very typical synapomorphy of Diamphidicus — ^the short M fork — is absent in H. setifera. Finally, a synapomorphy of the set of genera presently included in the Rhegmoclematini, the bottle-shaped aedeagus (with a large base and a distal bottle-neck), shared even by Diamphidicus, is absent in Holoclema. The representation in a cladogram of the set of features discussed above would result in a 3 -stemmed polytomy at the base of the Scatopsinae, with Holoclema, Rhegmocle- matini and Scatopsini+ as terminal taxa. Three other features may give some light to this problem. On the one hand, CUA2 is rather sigmoid in shape and there are no cerci, features apomorphic in all Rhegmoclematini. On the other hand, the male terminaha dorsally shows a pair of spiracles. The presence of spiracle pair Vm on the male terminaha is a plesiomorphic condition absent in ah other known Scatopsinae (known to occur only in the Aspistinae and Psectrosciarinae within the Scatopsidae). This sug- gests a sister-group relationship between Scatopse setifera Edwards and ah remaining Scatopsinae. The female terminaha gives some additional features that corroborate some of the previous inferences. The absence of a pair of gonapophysis VIQ laterahy to a mesal projection of stemite Vm — an apomorphic feature shared by the Rhegmoclematina and the Parascatopsina — ^is absent in Holoclema. This apomorphic condition of the female terminaha, however, is absent in Edwards’ species. The pair of lobes dorso-posteriorly on the terminaha, derived from tergite X (referred to by Cook as the cerci), is not fused to tergite VIII + IX, and again excludes the species from the Rhegmoclematina or from Diamphidicus (see Cook, 1971), which apomorphically possess this feature. The large, plesiomorphic tergite VIII + IX, on the other hand, excludes it from the Parascatopsina. 1996 EDWARDS’ TYPES OF NEOTROPICAL SCATOPSIDAE 45 Figs. 21-22. Pararhexosa tubifera (Edwards), n.comb. Female holotype. 21. Terminalia, ventral view; 22. Terminalia, dorsal view. These plesiomorphies of the female terminalia presented by H. setifera again suggest a basal placement somewhere at the base of the Scatopsinae. The position assumed here is that a new genus must be erected for Scatopse setifera Edwards and that it should be provisionally placed in the Rhegmoclematini. This would mean that the sigmoid CuA2 and the lost of cerci are truly synapomorphies uniting Holoclema and the remainder of the tribe. Its position in the Rhegmoclematini, hence, would be as the sister group to the remaining genera of the tribe. The spiracles, con- sequently, would have been lost twice in the Scatopsinae — in the Diamphidicina^ and in the Scatopsini^. Maybe more extensive studies of this species and the discovery of new species in the genus may help to better estabhsh its placement in the system of the family. Swammerdamellini Pararhexosa tubifera (Edwards), new combination Figs. 21-22 Scatopse tubifera Edwards, 1930:95, pi. DC, Figs. 6, 8 (wing, male terminalia). Type- locality: Chile, Llanquihue, Casa Pangue; Cook, 1967:2 (catalogue). The holotype of this species has six levels of labels: (1) a plastic strip with the specimen double mounted; (2) a label with “Casa Pangue/4-10.xii.l926” printed; (3) a plastic strip with the terminalia mounted; (4) a red, round label with “Type” printed; 46 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) (5) ''Scatopse tubifera Edw.” handwritten and “F. W. Edwards/det. 1930” printed; (6) S. CHILE/Llanquihue/E «& M. Edwards/B.M. 1927-63“. Contrary from what is in the description, the holotype of this species is a female, not a male. Edwards’ (1930, pi. DC, Fig. 8) drawing of the terminaha is a dorsal view, redrawn here to include the anterior part (Figs. 21-22). The maxillary palpus is long, reniform and there are setae on the front ventrally to the eye-bridge; the oceUi are adjacent to the eye-bridge. The tarsi are brown. The spiracular sclerite is small; there are 5 subspiracular setae, 2-30 anepistemals on the dorsal half of the sclerite, apparently no katepistemals and no merals, 3-5 mesepimerals. Abdominal stemites 2-6 are nor- mally sclerotized. The placement of this species also deserves some discussion. There is no doubt about its inclusion in the Scatopsinae. The large, reniform maxillary palpus would hardly allow any other placement than in the Swammerdamellini. The fact that abdominal stemites 2-6 are normally sclerotized shows that it certainly does not fit in the higher genera of the tribe, as Rhexosa, Abrhexosa, Quateiella and Hawomersleya. The small spiracular sclerite, however, indicates still a more basal position, since all these genera, as well as Coboldia, Swammerdamella, and Akorhexoza do possess elongated, large spiracular scle- rites. The monophyly of the genus Pararhexosa Freeman is still not well demonstrated. However, the species placed in this genus (some of which would fit in the generic diagnosis of Rhexosa) show a number of plesiomorphies that show the derived condition in all remaining genera of the Swammerdamellini. ACKNOWLEDGMENTS We must acknowledge the facilities given by Dr. Brian Pitkin at the British Museum for the examination of the types. This work was partially supported by FAPESP 94/2371-3 and CNPq 30253 1/88-2/ZO research grants. We also acknowledge a careful correction of the English version by an anonymous referee. LITERATURE CITED Amorim, D. S. 1982a. Classifica^ao por seqiiencia9ao: Uma proposta para a denomina9ao dos ramos retardados. Rev. Bras. Zool. 1(1): 1-9. Amorim, D. S. 1982b. Sistematica filogenetica dos Scatopsidae. Masters Thesis, Universidade de Sao Paulo, Sao Paulo, v + 173 pp + 74 pi. Amorim, D. S. 1994. A new suprageneric classification of the Scatopsidae (Diptera: Psychodo- morpha). Iheringia, Sen Zool. (77): 107-1 12. Cook, E. E 1955. A contribution toward a monograph of family Scatopsidae (Diptera). 2. The genera Rhegmoclemina Enderlein, Parascatopse n.g., and new species of Rhegmoclema. Ann. Entomol. Soc. Am. 48:351-364. Cook, E. E 1956a. A contribution toward a monograph of family Scatopsidae (Diptera). 3. The genus Rhexoza Enderlein. Ann. Entomol. Soc. Am. 49:1-12. Cook, E. E 1965. A contribution toward a monograph of family Scatopsidae (Diptera). 8. The genus Anapausis. Ann. Entomol. Soc. Am. 58:7-18. Cook, E. E 1967. 22. Family Scatopsidae, pp. 1-9 in: PAPAVERO, N. (ed.), A catalogue of the Diptera of the Americas south of the United States. Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo. Cook, E. E 1971. The Australian Scatopsidae (Diptera). Austr. J. Zool., Suppl., 8:1-90. Cook, E. E 1974. A synopsis of the Scatopsidae of the Palaearctic. 3. The Scatopsini. J. Nat. Hist. 8(1):61-100. 1996 EDWARDS’ TYPES OE NEOTROPICAL SCATOPSIDAE 47 Cook, E. E 1976. A new genus and five new species of Scatopsidae (Diptera) from New Guinea and Australia. J. Austr. Entomol. Soc. 15:447-452. Edwards, E W. 1930. Scatopsidae, p. 88-98, pis. 9-10 in: Diptera of Patagonia and South Chile based mainly on the material in the British Museum (Natural Elistory). V. 2, fasc. 3. British Museum (Natural History), London. Enderlein, G. 1912. Zur Kenntnis der Zygophthalmen. Uber die Gruppierung der Sciariden und Scatopsiden. Zool. Anz. 40:261-282. Freeman, P. 1985. Scatopsidae, pp. 20-48, 54-74 in: Freeman, P & R. P Lane, Bibionid and scatopsid flies — Diptera Bibionidae and Scatopsidae. Handbk. Ident. Brit. Insects 9(7). Royal Entomological Society of London, London. Received 23 April 1996; accepted 11 September 1996. J. New York Entomol. Soc. 104(l-2):48-6l, 1996 NEW TRIBAL PLACEMENT OF THE GENUS COSCINOCEPHALUS PRELL, 1936, WITH DESCRIPTION OF THE LARVA, PUPA AND ADULT OF A NEW SPECIES FROM MEXICO (COLEOPTERA: SCARABAEOIDEA; DYNASTINAE) Miguel A. Moron and Brett C. Ratclifee ‘Departamento de Biosistematica de Insectos, Institute de Ecologia, A.C. A.R 63 Xalapa, Veracruz 91000 Mexico ^Systematics Research Collections, W436 Nebraska Hall University of Nebraska, Lincoln, NE 68588-0514, U.S.A. Abstract. — A discussion is presented suggesting the transfer of Coscinocephalus from the Cyclocephalini to the Pentodontini. Coscinocephalus tepehuanus is described as new, based on immature and adult forms, from Durango state, Mexico. A key to the third-stage larvae of nine genera of American Pentodontini is presented. Finally, a brief commentary is given about life history information. Resumen. — Se presenta una discusion para justificar la transferencia del genero Coscinoce- phalus de la tribu Cyclocephalini a la tribu Pentodontini. Se describe Coscinocephalus tepe- huanus n. sp. con base en 6 adultos, asi como su larva de tercer estadio y la pupa, recolectados en un bosque de pinos y encinos del estado de Durango, Mexico. Se incluyen: una clave para separar las dos especies conocidas del genero, una clave para las larvas de tercer estadio de 9 generos de Pentodontini americanos y un comentario breve sobre los habitos de estas especies. Some genera of American Dynastinae have restricted geographical ranges and/or ecological tolerances, such as Aphonides Rivers, Pentodina Endrodi, Coscinoce- phalus Prell, Barutus Ratcliffe, Tehuacania Endrodi, Gillaspytes Howden, Hiekei- anus Endrodi, Neoryctes Arrow, Surutoides Endrodi, Indieraligus Dechambre, and Endroedianibe Chalumeau. Some of these genera are also monotypic, with insular habitats or extreme vicariant relatives and are usually very rare in collections. It is possible that most of these genera represent relictual elements that may provide useful data for phyletic studies of the subfamily. Recent collecting trips (both in the field and in collections) have given us new information on the controversial genus Coscinocephalus. Thus, the purposes of this paper are: (1) to justify the tribal transfer of Coscinocephalus from Cyclocephalini to Pentodontini; (2) to describe a new species from Mexico; (3) to describe the third- stage larva and pupa of the new species; (4) to update the known distribution of the genus; (5) to discuss the habits of the species; and (6) to provide a key to the known third-stage larvae of American Pentodontini. Technical terms used are those of Ritch- er (1966), Moron (1987) and Costa et al. (1988). Anoplocephalus cribrifrons was described by Schaeffer (1906) but the genus was preoccupied by Anoplocephalus Henrick 1895 (Platyhelminthes: Cestoda). Prell (1936) provided the replacement name Coscinocephalus. Schaeffer indicated the dif- ficulty of placing the genus in a tribe using the then present classification. He stated 1996 THE GENUS COSCINOCEPHALUS 49 that “this genus is intermediate between Lacordaire’s ‘Cyclocephalides’ and ‘Or- yctides’.” “Pentodontidae” was originally proposed by Burmeister (1847), but it was seem- ingly not widely used in elassifications even though Bates (1888) adhered to it in the Biologia. It is easy to understand Schaeffer’s indecision (especially if he was unfamiliar with Burmeister’s work) when he had only the old descriptions of the “Cyclocephalides” or “Oryctides” by which to place the new genus, and neither one seemed to properly accommodate Anoplocephalus. It was not until Casey (1915), nine years after the description of Anoplocephalus, that Pentodontini made its way into the primary North American literature. The tribe again fell into obscurity when it was not used in Leng’s (1920) catalog of North American Coleoptera, Arrow’s (1937) Coleopterorum Catalogus, Blackwelder’s (1944) checklist of the Coleoptera of Latin America, or Arnett’s (1968) beetles of the United States. Endrodi (1969, 1985) reinstated the use of Pentodontini and pro- vided the first modem definition of it. The definition, however, leaves a lot to be desired because monophyly was not addressed; some concerns remain that the Pen- todontini (as currently comprised) may be paraphyletic. Adult pentodontines are distinguished by the presence of tubercles, carinae, or a fovea on the head and/or pronotum; mandibles on the lateral margin with or without teeth; propygidium with or without a stridulatory stmcture; protibiae usually triden- tate, and apex of the metatibiae usually tmncate and margined with short, spine-like setae. Dimorphism between the males and females is slight. The larvae are characterized by 1-3 dorsal sensory spots on the last antennal segment; the molar area of the left mandible is dentate or, if lacking teeth, then the stridulatory teeth of the maxilla have acute projections directed distally; and the raster has one or more rows of palidia or, if lacking the palidia, then the chaetoparia of the epipharynx has few setae. Some adults in the Pentodontini are similar to others in the Oryctini. The two tribes (as currently stmctured) may generally be separated by the presence of a tmncate or nearly smooth apical rim on the apex of the posterior tibia in the pen- todontines and by a toothed (including apical tooth) or strongly crenulate rim in the oryctines. As opposed to the Cyclocephalini, all pentodontines have at least some head or pronotal armature in the form of carinae, tubercles, or even horns. It appears that Casey (1915) was the first to definitively place the then Anoplo- cephalus in the Cyclocephalini. Endrodi (1969, 1985) maintained its position in the Cyclocephalini based on the absence of head and pronotal armature and the presence of slender legs. In this regard, however, it should be noted that both species of Coscinocephalus possess as much of a developed boss, or tubercle, on the frons as in species of Orizabus, Pentodina, or Euetheola in the Pentodontini. The slender, long legs seem to be a variable character within tribes. In this case, we believe it is eorrelated with the arboreal habits of Coscinocephalus species. By contrast, most members of the Pentodontini have soil burrowing habits and have stouter (though not tmly fossorial) legs. We believe that Coscinocephalus is more appropriately placed in the Pentodontini for the following reasons: (1) the head is “armed”; (2) the mandibles, labmm, and maxilla are all relatively simple or small, and the labium is strongly constricted apically as in Orizabus (but unlike the general state found in the Cyclocephalini). 50 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) See Table 1 for further comparisons; (3) the clypeus is strongly bilobed and reflexed as in Orizabus but unlike any other Cyclocephalini; (4) the form of the parameres is more elongated, slender, and setose as in other Pentodontini but especially as in Orizabus; (5) the characters of the mouthparts in the larvae of Coscinocephalus, while showing no clear affinity (Table 2), are not like those of Cyclocephalini. Our study of both adult and larval features leads us to conclude that Coscinoce- phalus is not a cyclocephaline. In most respects the adults seem to be more closely related to Orizabus in the Pentodontini while the larvae share some affinities with larvae of Xyloryctes in the Oryctini. Even though we have some reservations about the monophyly of each of these tribes, we here place Coscinocephalus in the tribe Pentodontini. We have thus come full circle to Schaeffer’s (1906) original feeling that “this genus is intermediate between Lacordaire’s ‘Cyclocephalides’ and ‘Oryctides’.” The Pentodontini, a concept unknown or unrecognized by Schaeffer, is the intermediate between the Cyclocephalini and the Oryctini, and it is here that we place Coscino- cephalus. Coscinocephalus tepehuanus Moron and Ratcliffe, new species (Figs. 1-8) Type series: Holotype labeled “Mexico: Durango, Res. Biosfera ‘La Michiha,’ alt. 2,400 m, R. Terron col., pino-encino, ex larva, 24-X-87, adulto lO-IV-88, suelo cerca raices Quercus durifolia.'' Allotype labeled “Mexico: Durango, Suchil, Piedra Her- rada. Res. Biosfero ‘La Michiha,’ alt. 2,400 m, 16-VIII-79, bosque Quercus-Pinus, sobre suelo, M.A. Moron col.” Four paratypes with the following data: as holotype except without date of adult eclosion (1 male), as allotype except lO-VII-86 and “caminando en suelo, despues de Iluvia, 17 hrs. R.A. Terron Sierra col.” (2 females); “Mexico: Durango, Suchil, Res. Biosfera ‘La Michiha,’ Rcho. de la Pena, 26-VII- 78, R. Halffter col., luz, bosque Pinus-Quercus, alt. 2,400 m” (1 male). The pupal and third instar exuviae are also present for the holotype and one male paratype. Holotype and allotype deposited in the Miguel A. Moron collection (at the Insti- tute de Ecologia, Xalapa); two paratypes (male and female) deposited at the Institute de Biologia, UNAM collection (Mexico City); remaining male and female paratypes in the B.C. Ratcliffe collection (Lincoln). Holotype: Male. Length 21.5 nun from apex of clypeus to apex of elytra; width across humeri 11.2 mm. Color uniformly light reddish brown, moderately shining. Head: Clypeus with sides sinuate, converging to broad, bilobed apex; sides and apex strongly reflexed; surface densely rugopunctate. Vertex with large, oblong to trans- verse punctures. Low, rounded boss on frons between eyes. Interocular width equals 3.8 transverse eye diameters. Eye canthus with ventrolateral row of long, stout setae projecting laterally. Antenna with 10 segments, club subequal in length to segments 2-7. Mandibles (Fig. 1) small, not visible from above, lacking teeth, lateral edge with row of large setae, molar area small. Labium (Fig. 2) strongly contracted to small, narrow, feebly emarginate apex; lateral edges with stout setae (as long or longer than palps); ventral surface in basal half strongly and broadly depressed. Labrum (Fig. 3) small, subtriangular. Maxilla (Fig. 4) with small, subtriangular, se- Table 1 . General structure of adult mouthparts in 7 species of Dynastinae. 1996 THE GENUS COSCINOCEPHALUS 51 O QJ H O T3 0 73 Xi X X X -D D X) ~o o u td 3 ;-l . ^ _o 0 _ o _ _o o _o O _o T3 O ^ o u 3 •r1 ^ o B •n ^ O C g 'C B c g 'C B O 3 ’C 3 B o c o c c c c o C "" < < < < < < < 52 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) ca C Q o £ "o X a II 'o .t: < 'C li S 6 I S HH cd > s T3 ^ ^ S3 T3 l-H 4-3 (U cd OX) 6 > cd O O & cd T3 x: O C/l C T3 ‘o a C cd T3 (U T3 u (U T3 C :3 & cd td u c td o c c 'o a o x: 2 2 1/3 w -S c c (U c C u (/3 (U (U C/3 D C/) (/) jO -O X) cd a cd cd O 1996 THE GENUS COSCINOCEPHALUS 53 Figs. 1-7. Coscinocephalus tepehuanus n.sp. holotype S 1) Right mandible, dorsal view. 2) Labium, ventral view. 3) Labrum, dorsal view. 4) Right maxilla, dorsal view. 5) Parameres, caudal view. 6) Genital capsule, lateral view. Fig. 7) Genital plates (allotype) of C. tepehuanus, ventral view. Scale line = 1 mm. Most of mouthpcut setae have been removed for clarity. lose galea fused to small, unarmed lacinia; stipes with groove laterally for reception of retracted maxillary palp; palpus with 4 segments, 1st segment short, 2nd segment about 4 times length of first, 3rd segment 3 times length of first, 4th segment a little shorter than 2nd and 3rd segments combined and with longitudinally depressed sen- sory area on dorso-lateral surface extending from near base to just past middle of segment (over half length of segment). Pronotum: Surface punctate, extremely finely shagreened between punctures; punctures moderate in density and size on posterior half, larger and denser in anterior half and along sides where some become rugo- punctate. Sides evenly rounded, widest at middle. Sides and base with marginal bead. Scutellum minutely shagreened, lacking punctures. Elytra: Surface with 6 rows of punctate, furrowed striae between suture and humeral umbone and 4 distinct rows 54 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) of punctate striae on sides; punctures of striae moderate in size, subequal in size to those in posterior angle of pronotum, weakly ocellate. Intervals weakly convex, minutely shagreened, with irregularly spaced, sparse micropunctures. Pygidium: Pre- pygidium densely punctate. Pygidium regularly convex, surface with sparse, mod- erately sized punctures on disc, extreme base and angles weakly rugopunctate. Ven- ter: Thoracic stemites with moderately dense, long, fulvous setae. Abdominal ster- nites each with transverse row of setae at about middle. Prostemal peg long with blunt apex covered by long setae. Legs: Foretibiae tridentate, teeth subequally spaced. Foreclaws slender, simple. Posterior tibiae with 8 small denticles at apex. All femora with 4 rows of setae (1 dorsal, 1 ventral, 1 on each margin). All tibiae each with 2 rows of setae on dorsal surface. Tarsi subequal in length to respective tibiae. Parameres (Figs. 5-6): Slender, symmetrical, with small lateral tooth just past middle of shaft. Venter with long setae in apical half on ventral aspect. Allotype: Length 24.0 mm from apex of clypeus to apex of elytra; width across humeri 11.3 mm. As holotype except in the following respects: Color dark reddish brown. Head: Surface sculpturing a little more coarse. Boss on top of head stronger, a little higher. Apical segment of maxillary palpus with sensory depression less than half length of segment. Pygidium: Surface only weakly convex. Legs: Tarsi shorter than respective tibiae. Genital plates (Fig. 7): Ventral plates rounded, rugopunctate, covered with many long, stout setae. Variation: Males (2 paratypes) (Fig. 8): Length 22.5-23.0 mm; width across humeri 11.5-12.0 mm. Color varies from light to dark reddish brown. Boss on frons a little stronger in both specimens than in holotype, elytral striae more furrowed. Females (2 paratypes): Length 24.0 mm; width across humeri 12.0 mm in both specimens. Both female paratypes are similar to the allotype. Remarks: Coscinocephalus tepehuanus is most readily separated from C. cribri- frons by the characters listed in the key. The differences observed between these two species are indicative of recent separation and isolation of one species in the mountains of southeastern Arizona, Sonora, and Chihuahua from the much more southerly species in southeastern Durango. This scenario correlates well with the Pleistocene climatic events that permitted elevational lowering of vegetation. One result of this was that taxa living in the pine/oak associations of the Sierra Madre Occidental in Mexico were able to expand northward to the discontinuous mountains of southern Arizona. Subsequent warming and aridity reestablished the higher ele- vational gradient supporting pine/oak forests, which effectively created mountain islands harboring, at higher elevations, pine/oak forest (with its associated fauna) in a surrounding sea of lower elevation desert. See Jameson (1990) and Ratcliffe and Deloya (1992) for further information and references on climatic change. We suggest that the ancestors of both species of Coscinocephalus dispersed and subsequently became isolated from one another in this way. Morphological diver- gence is not yet substantial between the two species due to the relatively recent isolation event. Nevertheless, differences in body structure are already evident, and geographical (hence, reproductive) isolation is now complete. The limited number of specimens of Coscinocephalus collected in Chihuahua and Sonora are conspecific with the Arizona populations even though there is an absence of any mountains high enough to support pine/oak forests between those in southern Arizona and the north end of the Sierra Madre (about 31°N) in Mexico. Deep can- 1996 THE GENUS COSCINOCEPHALUS 55 Fig. 8. Habitus of Coscinocephalus tepehuanus n. sp. paratype 6 . Scale line = 10 mm. yons formed by the higher tributaries of Rio del Fuerte and Rio Mezquital (22°- 27°N) probably restrict the distribution of C. tepehuanus to the southern Sierra de Tepehuanes, Sierra de Durango, or only to the Sierra de Michis (Fig. 21). We predict that C. tepehuanus should be found in other areas of southern Durango because suitable habitat exists (dry pine/oak forest). A lack of collecting accounts for the absence of additional specimens of this new species. Etymology: Derived from tepehuan, meaning “the owner of the mountains” (Si- meon, 1988). Tepehuans are an ancient, native people of the Nahua language group who still live in some parts of the mountains of Durango. Third-stage larva: Head: Maximum width of head capsule 7 mm. Surface of cra- nium deeply and sparsely punctate, reddish brown. Frons (Fig. 9) with short, fine, sparse setae; each anterior angle of frons with 1 seta; remaining cranial surface with I epicranial seta, 1 dorso-epicranial seta, 1 paraocellar seta and 2 supraocellar setae, and some short, fine, sparse setae on each side. Clypeus with 4 lateral setae. Labrum ovate, slightly asynunetrical, with 2 central setae and 2 lateral setae on each side. Ocelli present, not pigmented. Epipharynx (Fig. 14) with haptomeral process entire, very prominent and sclerotized, without posterior setae; chaetoparia with few sen- silla; right chaetoparia with 50-60 spinelike setae; left chaetoparia with 40-45 spi- nelike setae; right acanthoparia with 6 short, spinelike setae; left acanthoparia with II short, curvate setae; right gynmoparia wide, without plegmatia; left gynmoparia 56 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Figs. 9-19. Coscinocephalus tepehuanus n.sp. third-stage larva. 9) Frontal view of cranium. 10) Right mandible, mesad view. 11) Left mandible, ventral view. 12) Left maxilla, dorsal view. 13) Stridulatory area of maxilla. 14) Epipharynx. 15) Labium-hypopharynx. 16) Last antennal segment. 17) III left abdominal spiracle. 18) Form of holes in respiratory plate. 19) Claw of left posterior leg. Fig. 20. C. tepehuanus pupa, II left abdominal spiracle. Scale lines = 1 mm. narrowed; acroparia with 4-5 setae; pedium wide; sense cone with 4 sensilla; laeo- torma curved toward center of pedium; mesal portion of dexiotormae expanded and fused with the sclerotized plate. Scissorial area of right mandible (Fig. 10) with 2 teeth before scissorial notch and with rounded projection on inner margin; molar area with 3 irregular lobes. Scissorial area of left mandible (Fig. 11) with entire cutting blade before scissorial notch; inner margin with 2 rounded, small projections; molar area with well developed distal lobe (Ml); acia acute; brustia well developed; ventral stridulatory area extended, with 35-38 transverse ridges; mesad of stridula- tory area is a puncture with 3 stout setae. Galea with well developed, conical, sharply 1996 THE GENUS COSCINOCEPHALUS 57 pointed uncus (Fig. 12); lacinia with 3 terminal unci fused at bases, the inner slightly shorter than others. Maxillary stridulatory area with 8-9 truncate-rounded, small teeth and with a rounded anterior process (Fig. 13). Hypopharyngeal sclerome asym- metrical, strongly produced on right side into rounded process (Fig. 15). Dorsal surface of last antennal segment with 2 oval, small, sensory spots (Fig. 16). Thorax: Thoracic spiracles 0.72 mm long and 0.91 mm wide; respiratory plate reddish brown, shaped as a closed “C”; lobes of respiratory plate nearly approximate; spiracular bulla prominent, rounded. Pronotum with well marked, reddish-orange, irregular formed, lateral scleromes with 2 slender fine setae. Dorsa of meso- and metathorax without setae. All legs with similar tarsal claws, claws pointed, each with 1 internal and 1 external basal setae (Fig. 19). Abdomen: Abdonfinal spiracles 0.70 mm long and 0.89 mm wide. Respiratory plate reddish brown, regularly shaped as a closed “C”; lobes of respiratory plate closely approximate (Fig. 17), with a maximum of 38-43 irregular shaped, ameboid “holes” along any diameter (Fig. 18); spiracular bulla pronunent, rounded. Chaetotaxy on dorsum of each segment as follows: I LS1(4): II LS2(2-10),SS4-5(10-16); III-V LS2-3(2-12),SS4-5(20-24); VI LSl(10),SS4-5(20-24); VII-VIII LS1(2); IX-X LS1(6). Venter of segments I-X each with 4LS. Raster without septula and palidia. Lower anal lip with 70-86 me- dium sized, spinelike setae. Upper anal lip with 26-40 short, spinelike setae and 10- 12 long, slender setae. Pupa: Body shape elongate, oval, stout, exarate. Color reddish orange. Head: Head bent sharply beneath thorax, mouthparts directed ventrally; antennae, mandibles, palpi, and compound eyes recognizable. Anterior border of clypeus slightly tumid. Thorax: Pronotum nearly transverse. Pterotecae closely appressed, curved ventrally around body, extending posteriorly between abdominal segments II-III. Protibiae with 3 rounded projections on exterior margin. Tarsomeres delineated. Abdomen: 9 visible segments. Spiracles of segment I ovate, slightly prominent, hidden by elytra and by oblique pleural fold; spiracles on segments II-IV ovate, annulate, sclerotized, prominent as rounded tubercles (Fig. 20); spiracles on segments V-VIII occluded, rosetiform, not prominent, sclerotized or pigmented. Tergal area of segments I-VI with 5 pair of deep, moderately sclerotized dioneiform organs {sensu Costa et al 1988). Last segment with 2 symmetrical, ventrolateral folds; without urogomphi; last stemite and genital ampulla hidden by folds. KEY TO THE SPECIES OF COSCINOCEPHALUS (ADULTS) 1. Color light to dark reddish brown. Elytral striae on disc with weakly ocellate punctures, punctures subequal in size to those in posterior angle of pronotum. Clypeus with lateral edge sinuate between apex and eye canthus. Southeastern Durango, Mexico C. tepehuanus Moron & Ratcliffe, n.sp. 1'. Color dark brown or black. Elytral striae on disc with distinctly ocellate punctures, punctures larger than those in posterior angle of pronotum. Clypeus with lateral edge not sinuate between apex and eye canthus. Southern Arizona, USA; Sonoran and Chi- huahuan Sierra Madre Occidental, Mexico (Fig. 21) C. cribrifrons (Schaeffer) Life history: Coscinocephalus cribrifrons is uncommonly encountered although it may be locally abundant. Part of the reason for its seeming rarity is the fact that adults have a brief period of frenetic activity only at dusk, and they are not strongly attracted to lights. On certain nights after the onset of the late summer rains, males 58 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Fig. 21. Known distribution of Coscinocephalus species. C. tepehuanus: (1) La Michilia, Durango. C. cribrifrons: MEXICO: Chihuahua: (2) Creel; (3) Madera; (4) Yepomera; USA: Arizona: (5) Santa Rita mountains; (6) Patagonia mountains; (7) Huachuca mountains; (8) Chiricahua mountains. Base map of northwest Mexico adapted from Garcia & Falcon (1974). form what are probably mating aggregations. They swarm to a certain tree (Ratcliffe has observed them on pine trees) where they rapidly run up and down the trunk with an audible scritching noise as their tarsi move over the bark. Warner (personal communication to Ratcliffe, November 1994) observed the males gradually ascend- ing to the top of the tree where, presumably, a female or females were releasing pheromones. Coscinocephalus tepehuanus probably has similar habits. We know very little of their life history. The slightly elongated legs seem to us to be an adaptation for their arboreal activities related to mating. We know little of the larvae. Two larvae of C. tepehuanus were found in the soil near the roots of a dead oak tree. It is interesting to note that the larvae of Xyloryctes jamaicensis (Drury) have been found beneath leaf litter in the soil feeding on roots or microrhizae (Ratcliffe 1991) as have the larvae for X. thestalus Bates (Moron 1976). The larvae of Coscinocephalus species may be feeding on the roots of live oak trees. 1996 THE GENUS COSCINOCEPHALUS 59 The mouthparts of the adults suggest a soft diet such as sap or perhaps pollen. There are no teeth on the mandibles and maxillae for cutting or chewing, and the labrum is reduced. If we eliminate the sap of pines (Pinus spp.) as having too many unpalatable secondary compounds, the next most likely sources of food in these habitats are Fagaceae and Erycaceae: oaks (Quercus spp.), madrones {Arbutus spp.), and “manizallas” {Arctostaphylos spp.). KEY TO THE AMERICAN GENERA OF PENTODONTINI BASED ON KNOWN THIRD-STAGE LARVAE (Modified from Ritcher 1966; Moron 1976; Moron & Deloya 1991; Lumaret 1991; Morelli 1992) 1. Raster with palidia and septula 2 1'. Raster without palidia and septula 3 2. Palidia monostichous, nearly parallel, each palidium consisting of 7-10 strongly com- pressed pali whose tips are slightly hooked. Last segment of antenna with 2 dorsal sensory spots. Ocelli present. Surface of cranium chestnut brown. Maximum width of head capsule 4.37 mm Euetheola 2'. Palidia polystichous, each palidium consisting of a patch of 5-7 irregular, longitudinal rows of sharp, cylindrical, spine-like setae. Palidia and septula extending across lower anal lip. Last segment of antenna with 2-4 dorsal sensory spots. Ocelli present. Surface of cranium reddish. Maximum width of head capsule 6.5 mm .... Ligyrus (Ligyrodes) 3. Inner margin of left mandible (between scissorial and molar areas) with a small tooth 4 3'. Inner margin of left mandible smooth between scissorial and molar areas 8 4. Last antennal segment with 2 dorsal sensory spots 5 4'. Last antennal segment with 5-8 dorsal sensory spots 7 5. Spiracles of abdominal segments I-VIII similar in size 6 5'. Spiracles of abdominal segments I and VIII slightly smaller than those of abdominal segments II-VII, which are similar in size. Surface of cranium light brown, slightly roughened, reticulate, without distinctive pits. Maximum width of head capsule 4.8 mm Ligyrus {s.str.) 6. Color of cranium yellowish-brown, surface with numerous small pores. Each side with 1 prominent, exterior frontal seta. Ocelli vague. Maximum width of head capsule 3 mm Oxygrylius 6'. Color of cranium reddish brown, surface deeply and sparsely punctate. Each side with- out exterior frontal setae. Ocelli present, well defined. Maximum width of head capsule 7 mm Coscinocephalus 7. Epicranium with a transverse, roughened carina extending to supraocellar areas. Last antennal segment with 5-6 dorsal sensory spots. Ocelli absent. Surface of cranium dark brown. Maximum width of head capsule 4.7 mm Neoryctes 7'. Epicranium smooth. Last antennal segment with 8 dorsal sensory spots. Ocelli small. Surface of cranium reddish brown. Maximum width of head capsule 7.2 mm Diloboderus 8. Last antennal segment with 1-2 dorsal sensory spots. Spiracles of abdominal segments I-IV similar in size; those of abdominal segments V-VIII progressively smaller pos- teriorly. Frons with dense covering of setae 9 8'. Last antennal segment with 4-5 dorsal sensory spots. Spiracles of abdominal segments I-VIII similar in size. Frons with sparse, short setae only in anterior half. Surface of cranium chestnut brown. Maximum width of head capsule 6.8 mm Philoscaptus 60 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) 9. Maximum width of head capsule 6-6.5 mm. Distance between lobes of respiratory plate of spiracles much less than dorsoventral diameter of bulla Orizabus 9' . Maximum width of head capsule 3. 8-5. 2 mm. Distance between lobes of respiratory plate of spiracles slightly to much less than dorsoventral diameter of bulla . . . Aphonus ACKNOWLEDGEMENTS Roberto Terron (UAM-Xochimilco, Mexico City) is thanked for collecting and donating the larvae of C. tepehuanus. We thank Carl Olson (University of Arizona), Scott McCleve (Douglas, AZ), and William Warner (Chandler, AZ) for locality records, and the assistance of two anon- ymous reviewers. Partial support of this article is provided by the project “Atlas de Coleoptera Scarabaeoidea de Mexico” (EB067/P 134/93 CONABIO-Mexico). LITERATURE CITED Arnett, R. H., Jr. 1968. The Beetles of the United States. American Ent. Inst., Ann Arbor. 1,112 pp. Arrow, G. 1937. Coleopterorum Catalogus, vol. 21, pars 156, Scarabaeidae: Dynastinae. W. Junk, Berlin. 124 pp. Bates, H. W. 1888. Biologia Centrali-Americana, Insecta, Coleoptera vol. 2, part 2, p. 296- 336. Blackwelder, R. E. 1944. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America, pt. 2. Bull. U.S. Natl. Mus. 185:189-341. Burmeister, H. 1847. Handbuch der Entomologie, band 5. TC.E Enslin, Berlin. 584 pp. Casey, T. L. 1915. A review of the American species of Rutelinae, Dynastinae and Cetoniinae. Mem. Coleop. 6:1-394. Costa, C., S. A. Vanin, S. A., and Casari-Chen. 1988. Larvas de Coleoptera do Brasil. Mus. Zool. Univ. Sao Paulo. 282 pp. (165 plates). Endrodi, S. 1969. Monographic der Dynastinae. 4. Tribus: Pentodontini. Ent. Abh. Mus. Tierk. 37:1-145. Endrodi, S. 1985. The Dynastinae of the World. Dr. W. Junk, Dordrecht. 800 pp. Garcia, E. and Z. Falcon. 1974. Nuevo Atlas Porrua de la Republica Mexicana. Ed. Porrua, Mexico. 197 pp. Jameson, M. L. 1990. Revision, phylogeny and biogeography of the genera Parabyrsopolis Ohaus and Viridimicus, new genus (Coleoptera: Scarabaeidae: Rutelinae). Coleop. Bull. 44:377-422. Leng, C. W. 1920. Catalogue of the Coleoptera of America, North of Mexico. John D. Sher- man, Mount Vernon, NY. 470 pp. Lumaret, J. P. 1991. Le genre Neoryctes Arrow, 1808: description de la larve et position taxonomique du genre (Coleoptera: Scarabaeoidea: Dynastidae: Pentodontini). Elytron 5:337-341. Morelli, E. 1992. Taxonomia de los estados immaduros edaficolas de coleopteros dinastidos (Tribus: Cyclocephalini, Oryctini y Phileurini) del Uruguay. Tesis de Maestria en Biol- ogia, Zoologia. Universidad de la Republica, Montevideo, Uruguay. 123 pp. Moron, M. A. 1976. Descripcion de las larvas de tres especies mexicanas de melolontinos (Coleoptera, Melolonthidae: Dynastinae y Rutelinae). An. Inst. Biol. Univ. Nac. Auton. Mexico (ser. Zool.) 47(2): 1 19-134. Moron, M. A. 1987. Los estados immaduros de Dynastes hyllus Chevrolat (Coleoptera: Mel- olonthidae, Dynastinae); con observaciones sobre su biologia y el crecimiento alometrico del imago. Eolia Entomol. Mex. 72:33-74. Moron, M. A. and C. Deloya. 1991. Los coleopteros lamelicornios de la Reserva de la Biosfera “La Michilfa,” Durango, Mexico. Folia Entomol. Mex. 81:209-283. 1996 THE GENUS COSCINOCEPHALUS 61 Prell, H. 1936. Beitrage zur Kenntnis der Dynastinae. Tiber Homonymierverhaltnisse der Na- men der Gattungun und Untergattungun. Entomol. Blatt 32:145-152. Ratcliffe, B. C. 1991. The scarab beetles of Nebraska. Bull. Univ. Nebraska St. Mus. 12:1- 333. Ratcliffe, B. C. and A. C. Deloya. 1992. The biogeography and phylogeny of Hologymnetis (Coleoptera: Scarabaeidae: Cetoniinae) with a revision of the genus. Coleop. Bull. 46: 161-202. Ritcher, R O. 1966. White Grubs and their Allies. Oregon St. Univ. Press, Corvallis. 219 pp. Schaeffer, C. 1906. On Bradycinetus and Bolboceras of North America, with notes on other Scarabaeidae. Trans. Am. Entomol. Soc. 32:249-260. Simeon, R. 1988. Diccionario de la Lengua Nahuatl o Mexicana. Siglo Veintiuno, Mexico, D.E 783 pp. Received 15 April 1995; accepted 16 October 1996. J. New York Entomol. Soc. 104(1 -2): 62-69, 1996 MORPHOLOGICAL CASTE DIFFERENCES IN NEOTROPICAL SWARM-FOUNDING POLISTINAE WASPS. \—PROTOPOLYBIA EXIGUA EXIGUA (HYMENOPTERA: VESPIDAE) Fernando B. Noll, Sidnei Mateus, and Ronaldo Zucchi Depaitamento de Biologia, Faculdade de Filosofia Ciencias e Letras de Ribeirao Preto. 14040-901 Ribeirao Preto (SP), Brasil; e-mail ifemnoll @ spider.usp.br Abstract. — A slight dimorphism between queens and, workers and intermediates was found in a colony of Protopolyhia exigua exigua collected in Pedregulho, southeastern Brazil. Mean sizes of all nine measured body parts of queens (n = 30) were statistically larger than those of workers and intermediates. Canonical discriminant analysis also showed some differentiation among them, with Mahalanobis’ intercaste distance (D^) 4.95 between queens and workers, 5.63 between queens and intermediates and 1.84 between intermediates and workers. Key words: Vespidae; Polistinae; Protopolyhia exigua exigua; caste differences; multivar- iate analysis. Among many fascinating aspects underlying colonial organization in social in- sects, caste related problems are surely outstanding. On such matters the most im- pressive aspect involves the diversity of caste patterns that, in turn, suggests the plasticity of evolutionary strategies leading to them. Neotropical swarm-founding Polistinae, in which polygyny is a rule, have remained less explored, mainly due to the difficulty of studying them continuously under both experimental and natural conditions. On the other hand, to some extent reliable sociological and biological information about these wasps can be obtained by analyzing a sample of wasps taken from a particular nest. This is especially true for caste differentiation and related phenomena. Caste differentiation in Polistinae is not pronounced (Richards, 1978). However, the Epiponini tribe has been morphometrically analyzed and, as pointed out by Rich- ards (1971, 1978; Richards and Richards, 1951), at least three caste differentiation stages are found: 1 -Conspicuous size and allometric differences present, with queens larger than workers in the absence of intermediates (sense Richards and Richards, 1951) (Agelaia spp: A. areata, Jeanne and Fagen, 1974; A. pallipes and A. multipicta, Simoes et al., in prep.; A. vicina, Noll et al., in prep.; Protonectarina sylveirae, Shima et al., 1996b); 2-Conspicuous dimorphism present, with queens smaller than workers and no intermediates present (Apoica flavissima, Shima et al., 1994; Polybia dimidiata, Maule-Rodrigues and Santos, 1974; Shima et al., 1996a); 3 -Morphological differences slight or indistinct, and intermediates present {Pseudopolybia vespiceps, Shima et al., in prep.). Evidently this group remains biometrically very little ex- plored. Most of the relevant papers (e.g. Richards and Richards, 1951; Richards, 1971, 1978) aimed primarily at caste distinction for taxonomical purposes, and so only characteristics supposedly more reliable for such aims were emphasized. 1996 CASTE DIFFERENCES IN PROTOPOLYBIA 63 The employment of a standardized methodology, as in multivariate analysis, has supported the establishment of patterns for the comparison of caste differentiation among the epiponine. Morphometric analysis on a Protopolybia exigua exigua col- ony are reported here showing that queens are slightly larger than workers and intermediates. MATERIAL AND METHODS The whole population, consisting of workers, intermediates and queens, was taken from a young colony of Protopolybia exigua exigua (de Saussure) collected in Ped- regulho (Southeastern Brazil) on Feb. 2, 1995. In order to detect morphological differences between castes the whole population was measured under a binocular microscope. The measured body parts were: head width (HW), minimum interorbital distance (IDm), gena width (GW), width of mesoscutum (MSW), alitrunk length (AL), length of gastral tergum I (TjL), basal height of Ti(TiBH), basal width of tergum II (T2BW), and partial length of the forewing (distance between the anterior edge of the first submarginal cell and the final margin of the marginal cell) (WL). The numerical data were statistically analyzed in relation to the ovarian and sper- mathecal states. The statistical analysis including the canonical discriminant analysis (CDA: Rao, 1973) were performed with the SAS Program Package for PC comput- ers, and Kruskal- Wallis One Way analysis of variance on ranks was used in order to detect caste differences for each character. If differences were detected, Dunn’s method for multiple comparison was performed using the program Sigma Stat for Windows version 1.0. RESULTS Nests and related aspects: P. exigua exigua occurs from NW South America (Co- lombia, Bolivia) to Southern Brazil (Richards, 1978) and, according to him: “the nests consist of a comb suspend from a leaf or branch by one or more peduncles and surrounded by an envelope in the side or bottom of which is an exit-hole; the envelope is attached to the sides of the combs. When there are additional combs they are built on the envelope of the previous one with a new envelope and exit hole. Generally, only one comb is found but one additional one is not rare.” The one-combed analysed nest (Fig. 1) had 307 cells in which 46 contained only eggs indicating the precocious state of the nest. It was attached by one principal peduncle and two auxiliaries ones. Concerning population size it was counted at 28 workers, 37 intermediates and 30 queens. Ovarian development and insemination: In the analyzed sample three kinds of ovary development were recognized (Fig. 2): type A, developed ovarioles bearing from two to several mature oocytes, type B, ovarioles with some oocytes at the begining of development and/or some in final phase of vitelogenesis and, type C, filamentous ovarioles bearing from no visible to slightly developed oocytes. Since only in the A type females the spermatheca contained sperm, females with ovaries A, B and C can be characterized as queens, intermediates and workers, respectively. As a probable consequence of the colony’s early stage, differences in the ovary condition between queens and intermediates were not clear and, in some cases, only insemination distinguished queens and intermediates. 64 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Fig. 1. Nest of Protopolybia exigua exigua. (Scale bar = 1.0cm). Caste differences in relation to morphometry: Among the mean relationships of the 9 analysed characters (Table 1), all of them show the comparisons between queen-worker and queen-intermediate significantly different. In contrast, intermedi- ate-worker comparisons were not significantly different {p < 0.05). The result of the canonical discriminant analysis based on 9 morphological char- Fig. 2. Kinds of ovary development found among the females of Protopolybia exigua ex- igua. A, developed ovaries found in inseminated females; B, developed ovaries found in un- inseminated females (intermediates) and C, undeveloped ovaries found in uninseminated fe- males (workers). (Scale bar = 1.0mm). Table 1. Means, CANj and CAN2 values and observed values of Dunn’s test for 9 characters used for discriminating castes of Protopolybia exigua exigua. 1996 CASTE DIFFERENCES IN PROTOPOLYBIA 65 ■^1 If a V -2 II 5 K 01 K 00 0 os CN so ON (N >n CO CN m q d d d 1 d 1 d 1 d 1 d d 1 CO ON VO CO CO so r- q q q q — 1 q q CO d 00 CN —1 in m so q q CO q m q q CO (N d CN (N width of pronotum; setae either brown or clear, stiff, like bristles 11 10. L with only 3 setae of different lengths [color pattern: DL and L light on both 1st and 4th (Table 1)] Hippodamia parenthesis, 1st instar — L with other setae in addition to 3 prominent ones, of various lengths [color pattern: DL and L light on both 1st and 4th lobes (Table 1)] Hippodamia parenthesis, 2nd instar 11. L with only 1 seta [color pattern: DL and L light on 1st; all lobes dark on 4th (Table 1)] Hippodamia convergens, 1st instar — L with 1 prominent long seta but several other shorter setae also present; [color pattern: DL and L light on 1st; on 4th segment variable, from some color on DL to both DL and L light (Table 1)] Hippodamia convergens, 2nd instar ACKNOWLEDGEMENTS My special thanks to Dr. Robert Pienkowski of the Department of Entomology, Virginia Polytechnic Institute and State University (VPI&SU), Blacksburg, Virginia, under whose guid- ance I undertook this project, and for whom I was privileged to work for two and a half years. He supplied the adult beetles, tested the key, and agreed to review this manuscript even though he was just beginning his retirement. I also want to thank Dr. R. D. Gordon of the Systematic Entomology Laboratory at Beltsville, Maryland, for his review of my first attempt at a key. His wise suggestions resulted in this final version, a much better key than the previous one. My appreciation goes to Scott Ludwig, currently a graduate student at the University of Georgia and formerly a graduate student in the Department of Entomology, VPI&SU, for supplying aphids for rearing the beetles. LITERATURE CITED Gage, J. H. 1920. The larvae of the Coccinellidae. Univ. of Dlinois Biological Monographs 6: 232-294. 88 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Gordon, R. D. and N. Vandenberg. 1991. Field guide to recently introduced species of Coc- cinellidae (Coleoptera) in North America, with a revised key to North American genera of Coccinellini. Proc. Entomol. Soc. Wash. 93(4);845-864. Gordon, R. D. and N. Vandenberg. 1993. Larval systematics of North American Cycloneda Crotch (Coleoptera; Coccinellidae). Entomol. Scand. 24(3):301-312. Palmer, M. A. 1914. Some notes on life history of ladybeetles. Annl. Entomol. Soc. America VII:2 13-240. Rees, B. E., D. M. Anderson, D. Bouk, and R. D. Gordon. 1994. Larval key to genera and selected species of North American Coccinellidae (Coleoptera). Proc. Entomol. Soc. Washington 96(3):387-412. Storch, R. H. 1970. Field recognition of the larvae of native Coccinellidae, common to the potato fields of Aroostook County. Maine Agric. Exp. Station Tech. Bull. 43, 16 pp. Strouhal, H. 1927. Die Larven der palaearktischen Coccinellini und Psylloborini (Coleopt.). Archiv fur Naturgeschicte. Abteilung A. 92:1-63. Van Emden, F. I. 1949. Larvae of British beetles. VII. (Coccinellidae). Entomol. Mon. Mag. 85:265-283. Received 24 June 1996; accepted 11 September 1996. J. New York Entomol. Soc. 104(l-2);89-94, 1996 SEASONAL OCCURRENCE OF BRACHYMERIA INTERMEDIA (HYMENOPTERA: CHALCIDIDAE), A PARASITOID OF THE GYPSY MOTH, LYMANTRIA DISPAR (LEPIDOPTERA: LYMANTRIIDAE), IN WESTERN MASSACHUSETTS V. Kerguelen and R. T. Carde^ Department of Entomology, Femald Hall, University of Massachusetts, Amherst, Massachusetts 01003 Abstract. — To document the life history of Brachymeria intermedia in the U.S., we monitored the presence of reproductively active wasps in the field using trap hosts over an 8-week time period in early summer. The overall level of recovery of wasps was low (8% peak parasitism on average), yet, all parasitism occurred within one week. Reproductively active B. intermedia seemed to be fairly synchronized with gypsy moth pupation, suggesting that they might be univoltine and monophagous. However, we cannot conclude unequivocally that B. intermedia is monophagous, because these wasps appear to disperse readily to other habitats where their behavior is unknown. Key words: Host finding. Brachymeria intermedia (Nees) was introduced from Europe to the United States as a potential biological control agent against the gypsy moth, Lymantria dispar L. It was released repeatedly from 1908 through 1963 (Hoy, 1976; Clausen, 1978), but its establishment was not confirmed until 1966 (Leonard, 1966). Dowden (1935) reported that in Europe B. intermedia completes one generation on the gypsy moth in late spring. A second generation develops on other lepidopteran hosts in late summer. However, owing to differences in climate between the Mediterranean areas where wasps were collected and New England, and therefore to differences in the phenology of the gypsy moth, it was suggested then that the life cycle of B. inter- media nfight differ if the wasp established in the United States. Howard and Fiske (1911) hypothesized that adult wasps lived long enough to allow a single generation life cycle with the gypsy moth as a sole host. Conversely, Dowden (1935) suggested that American B. intermedia might have a first generation on alternate hosts in the spring. Then, a second generation would develop on the gypsy moth (with a possible late third generation on alternate hosts.) Thirty years after the establishment of B. intermedia, the life cycle of the wasp in the United States is still unclear. Recoveries of wasps from species other than the gypsy moth have been rare (Prokopy, 1968; Leonard, 1975), yet many species are accepted as hosts in the laboratory (Dowden, 1935; Minot and Leonard, 1976; Roth- eray et al., 1984; Dindo, 1990; Drost and Carde, 1992a). Whether B. intermedia parasitizes other species before gypsy moth pupae are available in the field remains unknown. Although it was verified that females overwinter as adults (Waldvogel and ^ Current address: Department of Entomology, University of California, Riverside CA 92521. 90 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Brown, 1978; Schaefer, 1993), we do not know when wasps become active in the spring. In this study, we monitored the presence of reproductively active B. inter- media in the field, before, when, and after wild gypsy moth pupae were present. MATERIALS AND METHODS Study sites: The study was conducted in three sites, Ludlow, Amherst and Mont- ague, Massachusetts. These sites were selected for their high densities of gypsy moth the preceding years. The sites in Ludlow and Amherst were dominated by oaks (Quercus rubra L., Q. alba L. and Q. prinus L.). The site in Montague was a mixed stand of birch (Betula populifolia Marshall), oaks {Q. rubra, Q. ilicifolia Wangen- heim), pines (Pinus rigida Miller) and willows {Populus tremuloides Michaux). Monitoring: We monitored the presence of actively reproductive wasps by exposing trap hosts continuously in the field from June 2, to July 21, 1994 in Ludlow (8 weeks) and June 6 to July 21, 1994 in Amherst and Montague (7 weeks). Trap hosts are nonparasitized hosts reared in the laboratory, placed in the field and recovered after short intervals of time to be checked for parasitism (Van Driesche et al., 1991). At each site, one cage holding 10 two to five day-old male gypsy moth pupae, reared from eggs on artificial diet in the laboratory (Bell et al., 1981), was stapled on 10 trees, 3 to 5 meters apart (oak in Ludlow and Amherst and birch in Montague). Thus, 100 pupae were exposed at each site. Cages were made of 7 nun wire mesh and placed ca. 2 m above ground level on the south side of trees. Pupae were renewed weekly and subsequently reared in the laboratory at 24 ± 1°C, 60-70% relative humidity with a 16 L:8 D light cycle. They were checked daily for emergence of moths or B. intermedia. Deaths due to predation in the field and to unexplained causes were also recorded. Mortality rates were computed as the marginal rates of attack, i.e., the rates of mortality due to each mortality agent in the absence of other contemporaneous agents (Buonaccorsi and Elkinton, 1990; Elkinton et al., 1992). Mortality due to predators was computed as the observed death rate. Mortality due to B. intermedia and unexplained causes was computed as the marginal rates of attack for two contemporaneous indiscriminate parasitoids (with c = 0.5). RESULTS The outcome of the rearing of pupae exposed in Ludlow, Amherst and Montague is presented in Fig. la, lb and Ic, respectively. Overall, most pupae escaped para- sitism and survived to the adult stage at all three sites (69, 49 and 77% average adult emergence, respectively), with little variation over the course of the experiment. Parasitism by B. intermedia was low at all sites and occurred only during one week: 15% parasitism (12 wasps recovered) at Ludlow on week 3 (nud-June), 1% parasit- ism (1 wasp recovered) at Amherst on week 5 (early July) and 9% parasitism (8 wasps recovered) at Montague on week 5. Predators (beetles, hemipterans, and ants) had a low impact (8, 35 and 3% average predation, respectively), although a high predation rate (77% on average), mainly attributed to ants, was observed the last three weeks of study at Amherst. Larger predators including mice, were excluded by the cages. Mortality due to undermined factors averaged 24, 39 and 19%, re- spectively. 1996 SEASONAL OCCURRENCE OF BRACHYMERIA INTERMEDIA 91 b ^B.int □ Pred ^L.disp □ Other Week Fig. 1. Fate of gypsy moth pupae exposed weekly in the field from June 2 to July 21, 1994 at three study sites (a- Ludlow, b- Amherst and c- Montague, MA) and subsequently reared in the laboratory. (B.int: emergence of B. intermedia, Pred: killed by predators, L.disp: moth emergence. Other: unexplained death. Weekly total of pupae equals 100.) DISCUSSION B. intermedia is a parasitoid with the most impact in high density gypsy moth populations. The incidence of parasitism has been correlated with egg mass counts, larval and pupal densities of gypsy moth, and percentage of defoliation (Reardon, 1976; Ticehurst et al., 1978; Williams et al., 1993). Although densities of gypsy 92 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) moths were fairly high at the study sites the preceding years, density of pupae was low at all sites in 1994 because larvae were decimated by an epizootic of nuclear polyhedrosis virus and fungal pathogen Entomophaga maimaiga Humber, Schimazu and Soper. Therefore, none of the sites was defoliated during this study and few field pupae were available for the wasps to parasitize. Coincidentally, the number of wasps recovered in our experiment was low at all three sites. Considering the scarcity of pupae in the field, one might have expected that trap hosts would be particularly attractive to the wasps should they be present. If B. intermedia is monophagous on the gypsy moth, one might expect peak levels of parasitism to occur when the host population declines following an outbreak (delayed density dependence). However, Ticehurst et al. (1978) also observed a collapse of parasitism concurrent with the collapse of the host population. The low level of parasitism of trap hosts, suggests that either the small number of pupae was not sufficient to retain the wasps in the area, or that the wasps located sparsely distributed hosts with difficulty. Blumenthal et al. (1979) suggested that B. intermedia readily dispersed en masse from sparse infestation sites and colonized high density popu- lations. Although trap hosts were present in the field for eight or seven weeks, it is striking that wasps were recovered within a single week at each site. These hosts were parasitized approximately two weeks before peak pupation (gypsy moth development was almost two weeks advanced at Ludlow compared to the other two sites). Ap- parently, B. intermedia was present in the field for a limited period of time, somewhat synchronized with pupation of the gypsy moth. The sites of Amherst and Montague had been visited weekly from late spring to early summer to search for B. intermedia. The leaf litter, dead trees and crevices were examined within ca. 5,000 square meters around the study sites of Amherst and Montague as they nfight have sheltered ag- gregated overwintering wasps (Dowden, 1935; Waldvogel and Brown, 1978), but no wasps were located. This suggests that B. intermedia was not present on these sites early in the season and that the wasps arrived near the time of pupation of gypsy moth larvae. In the absence of sufficient numbers of pupae, B. intermedia might have emigrated elsewhere, as suggested by Blumenthal et al. (1979). Even when wasps did parasitize trap hosts, only a small fraction was' parasitized. While 12 pupae out of 100 were parasitized in Ludlow, 54 were evidently not, as adults emerged later. In Montague 8 pupae were parasitized and 78 were not. In Amherst one pupa was successfully parasitized and 79 were not. In addition, the pupae in a cage were never all parasitized. In Ludlow, 6 pupae were parasitized in two cages on neighboring trees. In Montague, the 8 parasitized pupae were found in 5 different cages. Female B. intermedia produce 5 offspring on average per day (Dowden, 1935; Barbosa et al., 1986), and can successfully parasitize up to 11 pupae in a row in the laboratory (Drost and Garde, 1992b). Therefore, a single wasp po- tentially could parasitize most of the pupae in a cage on a single visit. However, previous direct observations of the behavior of B. intermedia in the field (in 1993 in Montague, unpublished data) revealed that the wasps almost always visited only one pupa within a cluster. The present results corroborate these findings. Despite the small number of wasps recovered, our results indicate that the popu- lation of reproductively active B. intermedia was reasonably synchronized with the availability of gypsy moth pupae. Yet, the life history of B. intermedia in the field 1996 SEASONAL OCCURRENCE OF BRACHYMERIA INTERMEDIA 93 remains equivocal as the apparent synchrony with the gypsy moth population may indicate that the wasps either use the gypsy moth as a unique host or parasitize alternate hosts in other habitats. AC KNO WLEDGMENTS We thank Dr. J. S. Elkinton for valuable discussion. We thank Vic Mastro and G. Bemon of APHIS Methods Development Laboratory, Otis, MA, for supplying gypsy moth egg masses. This research was supported by a USDA National Research Initiative grant. LITERATURE CITED Barbosa, R, P. Martinat, and R. Bennet. 1986. Consequences of maternal age and host depri- vation on the production and development of Brachymeria intermedia (Nees) and the mortality of its gypsy moth (Lymantria dispar) host. J. Appl. Entomol. 101:215-223. Bell, R. A., C. D. Owens, M. Shapiro, and J. G. R. Tardif. 1981. Development of mass rearing technology. Pages 599-631 in: The gypsy moth: research toward integrated pest man- agement. C. C. Doane and M. L. McManus (eds.) — USDA, Techn. Bull. 1584. Blumenthal, E. M., R. A. Fusco, and R. C. Reardon. 1979. Augmentative release of two established parasite species to suppress populations of the gypsy moth. J. Econ. Entomol. 72:281-288. Buonaccorsi, J. P. and J. S. Elkinton. 1990. Estimation of contemporaneous mortality factors. Res. Popul. Ecol. 32:151-171. Clausen, C. P. 1978. Introduced parasites and predators of arthropod pests and weeds: a world review. USDA, Agr. Handbook. 480. Dindo, M. L. 1990. Comportemento di oviposizione di Brachymeria intermedia (Nees) (Hym. Chalcididae) in crisalidi di Galleria mellonella L. (Lep. Galleriidae). Boll. 1st. Entomol. G. Grand! Univ. Bologna. 45:109-119. Dowden, P. B. 1935. Brachymeria intermedia (Nees), a primary parasite and B. compsilurae (Cfwd.), a secondary parasite, of the gypsy moth. J. Agric. Res. 50:495-523. Drost Y. C. and R. T. Carde. 1992a. Host switching in Brachymeria intermedia (Hymenoptera: Chalcididae), a pupal endoparasitoid of Lymantria dispar (Lepidoptera: Lymantriidae). Environ. Entomol. 21:760-766. Drost, Y. C. and R. T. Carde. 1992b. Influence of host deprivation on egg load and oviposition behaviour of Brachymeria intermedia, a parasitoid of gypsy moth. Physiol. Entomol. 17: 230-234. Elkinton, J. S., J. P. Buonaccorsi, T. S. Bellows, Jr., and R. G. Van Driesche. 1992. Marginal attack rate, k-values and density dependence in the analysis of contemporaneous mor- tality factors. Res. Popul. Ecol. 34:29-44. Howard, L. O. and W. E Fiske. 1911. The importation into the United States of the parasites of the gypsy moth and the brown-tail moth: A report of progress, with some consider- ations of previous and concurrent efforts of this kind. USDA Bur. Entomol. Bull. 91: 240-252. Hoy, M. A. 1976. Establishment of gypsy moth parasitoids in North America: An evaluation of possible reasons for establishment or non-establishment. Pages 215-232 in: Perspec- tives in Forest Entomology. J. F. Anderson and H. K. Kaya (eds.). Academic Press Inc., New York. Leonard, D. E. 1966. Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae) established in North America. Entomol. News. 11 -.7.5-21 . Leonard, D. E. 1975. Parasitization of the spruce budworm, Choristoneura fumiferana (Clem- ens) (Lepidoptera: Tortricidae) by Brachymeria intermedia (Nees) (Hymenoptera: Chal- cididae). J. New York Entomol. Soc. 83:269-270. 94 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Minot, M. C. and D. E. Leonard. 1976. Host preference and development of the parasitoid Brachymeria intermedia in Lymantria dispar, Galleria mellonella, and Choristeoneura fumiferana. Environ. Entomol. 5:527-532. Prokopy, R. J. 1968. Parasites of the leaf rollers Archips argyrospilus and A. griseus in Con- necticut. J. Econ. Entomol. 61:348-352. Reardon, R. C. 1976. Parasite incidence and ecological relationships in field populations of gypsy moth larvae and pupae. Environ. Entomol. 5:981-987. Rotheray, G. E., P. Barbosa, and P. Martinat. 1984. Host influences on life history traits and oviposition behavior of Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae). En- viron. Entomol. 13:243-247. Schaefer, P. W. 1993. Overwintering aggregations of female Brachymeria intermedia (Hyme- noptera: Chalcididae). Entomol. News. 104:133-135. Ticehurst, M., R. A. Fusco, R. P. Kling, and J. Unger. 1978. Observations on parasites of gypsy moth in first cycle infestations in Pennsylvania from 1974-1977. Environ. Entomol. 7: 355-358. Van Driesche, R. G., T. S. Bellows, J. S. Elkinton, J. R. Gould, and D. N. Ferro. 1991. The meaning of percentage parasitism revisited: Solutions to the problem of accurately es- timating total losses from parasitism. Environ. Entomol. 20:1-7. Waldvogel, M. G. and M. W. Brown. 1978. An overwintering site of the gypsy moth parasite, Brachymeria intermedia. Environ. Entomol. 7:782. Williams, D. W, R. W. Fuester, W. W. Metterhouse, R. J. Balaam, R. H. Bullock, R. J. Chianese, and R. C. Reardon. 1993. Incidence and ecological relationships of pupal parasitism by Brachymeria intermedia in New Jersey populations of the gypsy moth. Entomophaga. 38:257-266. Received 13 May 1996; accepted 16 October 1996. J. New York Entomol. Soc. 104(1 -2): 95- 103, 1996 GROUND BEETLES OF NANTUCKET ISLAND, MASSACHUSETTS: 1995 (COLEOPTERA: CARABIDAE) Foster Forbes Purrington Department of Entomology, The Ohio State University, Columbus, Ohio Abstract. — A single season (1995) of ground beetle collecting on Nantucket Island, Massa- chusetts yielded 102 species in 18 tribes (Coleoptera: Carabidae). Fifty-seven species not pre- viously recorded from the island were taken, including five species not currently listed for Massachusetts. Globally rare habitats including Maritime Heath/Grasslands, Morainal Frost Bot- toms and Coastal Plain Ponds were intensively assayed. A high species turnover rate is implied by this preliminary analysis. The eastern United States has been surveyed for insects since the early nineteenth century. Throughout this region, the ground beetle fauna is generally very well known, but Nantucket Island, Martha’s Vineyard and the Elizabeth Islands off Mas- sachusetts south of Cape Cod in particular seem to have been especially neglected by entomologists, as measured by published research, due at least in part to their relative inaccessibility. Their isolation coupled with rarity of the maritime sandplain habitats developed there ironically makes these islands especially likely to contain unique assemblages of carabids, as well as species that may be rare elsewhere. Modem floristic studies on the islands present corresponding parallels (Peter W. Dunwiddie, pers. conun.). This report provides a checklist (Table 1) of 102 ground beetles that presently occur on Nantucket Island. Collections were made from early May to mid-September 1995, principally with pitfall traps containing propylene glycol-based preservative, using standard techniques. Voucher specimens are held at the Sandplains Bioreserve office of The Nature Conservancy, Vineyard Haven, Massachusetts. Several special island habitats, unique for their floristics, geology, hydrology and/ or microclimate, were singled out for intensive study, including the following sites, accounting for the large majority of ground beetle species collected in 1995: Miacomet Plains Maritime Heath/Grassland: near coastal sand dunes at southern end of Somerset Road; flat open terrain with scattered pitch pine, arrowwood copses and black cherry; also present: dwarf blueberry, huckleberry, Rubus, Rosa, Baptisia, Carex, trailing arbutus, bearberry, bracken fern, dense foliose lichen, and such old field serai forb types as milkweed, wood lily, thistle and Solidago. Morning fog, salt spray from the nearby south coast surf, lower ambient temperatures (relative to Nantucket town) and frequent sand-bearing onshore winds also define this habitat; an approximately 10 day lag in floral phenology compared with the town was ap- parent. Oak Barrens: xeric site west of Sankaty Head in the east central moors south of the Barnard Valley Road, dominated by thickets of mixed ilicifolialprinoides scmb oaks (Quercus). In late May 1995 a profound frost here killed much foliage and twigs. Other plants include Rubus, arrowwood, huckleberry, Carex and grasses. Table 1. Ground beetles (Coleoptera: Carabidae) collected on Nantucket Island in 1995. Tribes and genera in phylogenetic sequence (basal to derived), from Bousquet and Larochelle (1993); species listed alphabetically, noting European introductions. Ground Beetle Taxa Johnson (1930) 96 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(1-2) Bembidion confusum Hayward Bembidion constrictum (LeConte) Bembidion contractum Say Bembidion frontale (LeConte) 1996 CARABIDAE OF NANTUCKET 97 a X a o (D U Q w o 3 a 6 .§ a o o a a (D (U CQ m ^ o OJ M O w hJ a a *a gas S S P a a cd C/2 C C o cd C/2 -S -5 S ^ m pQ CQ m w pq clh ^ .a I i 0> cd PU PkH PU Oh C« t/3 (/) C/1 C/5 C/3 ■ cd HP (U w ^ ’5? g M S C c« (U 13 3 -td E £ a c/1 c/3 c/3 P3 :3 3 JH -C JP CJ U 26 mm, females > 29 mm); noticeably long-legged, length of profemur always longer than lateral length of pronotum by more than 1 mm; southeastern Brazil, Colombia C. volxemi (Montandon) 14b. Body shorter (males < 26 mm, females < 27 mm); not noticeably long-legged, length of profemur longer than lateral length of pronotum by up to 1 mm 15 15a. Parasterna of last abdominal segment posteriorly ending in large, thumblike processes (Fig. 30C); vertex usually without pronounced carina; posterolateral metasternal pro- cesses as in (Fig. 30B); Venezuela, the Guyanas, northern Brazil C. doesburgi De Carlo 15b. Parasterna of last abdominal segment posteriorly ending in digitate but not thumblike processes (Fig. 3 1C); vertex with pronounced carina; posterolateral metasternal pro- cesses as in (Fig. 3 IB); Colombia C. intermedia (Martin) Species Descriptions Curicta bilobata Kuitert Figs. 15, 32 Curicta bilobata Kuitert, 1949a;61-62; De Carlo, 1951:419. Redescription Measurements. Holotype male (no other specimens known): length, 19.2; profemoral length, 5.9; siphons, 12.5. Color. Light brown. Lighter mottling not apparent on thoracic or hemelytral dorsa. Abdominal dorsum reddish. Abdominal venter dark with four thin, lighter longitu- dinal bands on sterna 3 and 4 and a single broad band on parasterna 3-6. Protibia somewhat darker basally; no dark medial annulation evident. Protarsus absent on the one prothoracic leg present with specimen. Meso- and metathoracic legs apparently light brown. Structural characteristics. Body elongate (Fig. 15 A): length 5.5 X maximum width. Eye width 0.5 X interocular distance. Vertex broadly convex, domelike; without median longitudinal carina. Lobe of antennal segment 2 greater than 0.5 X length of 3. 146 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-J) Fig. 15. C. bilobata: A) dorsal habitus (5 mm); B) Male operculum (0.5 mm); C) Profemur, anterior view (0.5 mm); D) Distal fifth of profemur showing tubercles in sulcus, dorsal view (0.5 mm); E) Paramere (0.25 mm). 1996 THE GENUS CURICTA 147 Pronotum elongate; lateral length 1.5X posterior width. Lateral margins parallel anteriorly, divergent posteriorly. Transverse sulcus incomplete. Median longitudinal sulcus very shallow; obsolescent posteriorly. Median longitudinal ridges rounded; converging posterior to obsolescent median sulcus into single rounded ridge which continues to posterior margin of pronotum. Lateral sulci shallow. Lateral ridges rounded. Ridges and humeral lobes apparently without granulations. Posteroventral extensions of pronotum meet in ventral midline at junction of pro- and mesosterna (Fig. 2B). Scutellum width less than length. Trident pattern indistinct; transverse sections of lateral prongs slightly darker than rest of trident. Hemelytra widest just anterior to base of membranes; apparently no granulations present. Metathoracic wings fully developed. Plates of last abdominal tergum deformed by partially protruding phallus; no lon- gitudinal carinae or bands of hairs; apparently not tomentose. Prosternum in lateral view not swollen. Mesosternum with midline groove present; pair of narrow, parallel ridges on either side of the groove faintly apparent. Parasterna of last abdominal segment ending in small, slender digitate processes which do not extend posteriorly beyond posterior margin of the operculum. Male operculum. Fig. 15B. Procoxae 0.59 X profemoral length. Single truncate profemoral tooth anteroventral to profemoral sulcus (Fig. 15C); distance from base of femur to tip of anteroventral profemoral tooth 0.52 X total femoral length. Profemur moderately arched beyond profemoral teeth. Single row of tubercles in profemoral sulcus evident in distal half, becoming an irregular double row close to the tibia (Fig. 15D); no teeth in sulcus distally. Metafemora reaching onto proximal half of abdominal sternum 6 when extended posteriorly. Paramere, Fig. 15E (drawn from paramere mounted on plastic card beneath spec- imen; dissection of male genitalia not done in order to preserve condition of last abdominal tergal plates [see above]). Female genitalia not examined (known only from male holotype). Discussion. Kuitert diagnosed this species by the “shape of the anterior femur and by two large longitudinal plates on the dorsum of the last abdominal segment for which the species receives its name” (1949a: 62). The abdominal plate deformation, as noted above, is caused by a protruding phallus and therefore cannot be considered diagnostic. However, the single, truncate profemoral tooth is diagnostic. This species cannot be confused with the other species known to occur on Brazil’s coast, C. volxemi, doesburgi, and decarloi, because all three of these species have two pro- femoral teeth. In addition, volxemi and doesburgi have large thumblike parasternal processes while bilobata has small, slender processes. Finally, decarloi has a carina on the vertex, a dark, medial protibial annulus, and carinate pronotal ridges, while bilobata has none of these features. Distribution (Fig. 32). Brazil: Espirito Santo. Material Examined. HOLOTYPE, male: Espirito Santo, Brazil, leg. Fruhstorfer (no temporal data). Deposited in the Snow Entomological Museum, University of Kansas. 148 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Curicta bonaerensis (Berg) Figs. 16, 32 Helotenthes bonaerensis Berg, 1879a:72; 1879b: 194. Lectotype here designated. Nepa bonaerensis von Ferrari, 1888:191. Nepoidea falloui Martin, 1898:66-68. Syn. by Kirkaldy and Torre Bueno, 1909:202. Curicta boncerensis, Montandon, 1903:98-99. Curicta falloui, Montandon, 1903:98-99. Curicta bonaerensis, Kirkaldy and Torre Bueno, 1909:202; De Carlo, 1951:414- 415. Redescription Measurements. Males: length, 13.0-13.5; profemoral length, 3. 3-3. 7; siphons, 7.7- 8.4. Females: length, 13.2-16.0; profemoral length, 3. 4-3. 8; siphons, 6.2-9. 0. Color. Light brown. Lighter mottling not evident on thoracic or hemelytral dorsa. Abdominal dorsum brownish red. Abdominal venter dark, usually with four lighter, longitudinal bands on sterna 3-6; one such band per parasterna 3-6. Protibia with dark basal annulation, occasionally dark distally but no dark medial annulation pres- ent. Meso- and metathoracic legs light brown to golden. Structural Characteristics. Body elongate (Fig. 16A); length 4.2— 4.5 X maximum width. Eye width usually slightly greater than 0.5 X interocular distance. Vertex broadly convex; infrequently subcarinate medially. Lobe of antennal segment 2 usually up to 0.5 X length of 3. Pronotum subelongate; lateral length 1.1 X posterior width. Median longitudinal sulcus shallow, complete or obsolescent posteriorly. Transverse sulcus incomplete. Median longitudinal ridges rounded. Lateral sulci shallow. Lateral ridges rounded. Ridges and humeral lobes without granulations. Posteroventral extensions of pro- notum do not meet in ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width slightly less than length. Trident pattern usually indistinct; uni- form color. Hemelytra widest anterior to base of membranes; some granulations evident along hemelytral margins. Metathoracic wings fully developed. Last abdominal tergum variable: some specimens exhibit two narrow, postero- medial, parallel bands of hairs, while others are simply tomentose with no apparent bands. Prosternum in lateral view moderately swollen in posterior half. Mesosternum: midline groove present; longitudinal, parallel, tomentose bands often present, usually irregular and limited to posterior half. Metasternum tomentose; posterolateral pro- cesses robust (Fig. 16B). Parasterna of last abdominal segment ending in small, digitate processes which are subequal posteriorly to posterior margins of male and female opercula. Male operculum. Fig. 16C. Procoxae about 0.54X profemoral length. Single, pronounced profemoral tooth on the anteroventral margin of profemoral sulcus; distance from base of profemur to tip of anteroventral profemoral tooth about 0.59 X total femoral length. Profemur dra- matically arched beyond profemoral tooth (Fig. 16D). Profemoral sulcus curvilinear, convex posteriorly (Fig. 16E); no tubercles or sulcal teeth. 1996 THE GENUS CURICTA 149 Fig. 16. C. bonaerensis: A) dorsal habitus (5 mm); B) posterolateral metasternal processes, ventral view (0.5 mm); C) male operculum (0.5 mm); D) profemur, anterior view (1 mm); E) profemur, ventral view (1 mm); F) parameres (0.25 mm). 150 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-J) Metafemora reaching onto proximal half of abdominal sternum 6 when extended posteriorly. Parameres, Fig. 16F. Female genitalia: Gp2 arched dorsomedially (Fig. 13D). Discussion. The characters of the profemur readily separate this species from all other Curicta spp.: single elongate profemoral tooth; profemur dramatically arched beyond the profemoral tooth; and curvilinear profemoral sulcus. C. pelleranoi is similar to bonaerensis having a single profemoral tooth, a curvilinear sulcus, and similar parameres. However, pelleranoCs profemur is not so dramatically arched beyond the profemoral tooth. In addition, bonaerensis specimens have a moderately dilated prosternum in lateral view while pelleranoi specimens do not. Distribution (Fig. 32). Argentina: Buenos Aires, Entre Rios. Uruguay: Durazno. Material examined. Berg did not designate a type specimen for this species and it is unclear from his original description whether he was looking at a series or a single specimen. One of the specimens loaned from the Natural History Museum of the National University of La Plata has a determination label on it in Berg’s handwriting (Horn and Kahle, 1937, Volume 3, Table XXII, no. 9). I therefore am designating this specimen a lectotype. The type for Nepoidea falloui Martin bears a handwritten determination label with the species name, the word “type,” the name “J. Martin,” and the date “28 fe’r 1898.” It is clear from Martin’s original description that he was looking at a single specimen (“un exemplaire de la collection G. Fallou”, p. 66) therefore the specimen I have before me must be the holotype and I have so labeled it. LECTOTYPE, female: Helotenthes bonaerensis Berg. Argentina, Buenos Aries, leg. Guenther. No temporal data. Deposited in Natural History Museum of the National University of La Plata (MLP). HOLOTYPE, female: Nepoidea falloui Martin. Rio-Grande, leg. G. Fallou, “259-95.” No temporal data unless the three numbers listed are the date collected, e.g., 25 September 1895. Deposited in the Museum National D’Histoire Naturelle, Paris (MNHN). Additional material examined. 26 specimens collected in March (2), April ( 1 ), Sept. (1), Dec. (1) (the remaining 21 specimens without temporal data); deposited in JTPC, MACN, MLP, NMNH, MNHN, SEMC. ARGENTINA. Buenos Aires: Buenos Aires; Chascomas; Escobar; La Plata; Lujan; Rosas-F. C. Sud. Entre Rios: Lazo. URU- GUAY: Durazno: Castillo. Curicta pelleranoi De Carlo Figs. 17, 32 Curicta pelleranoi, De Carlo, 1951:415—416. Redescription Measurements. Male (1 specimen): length, 13.2; profemoral length, 3.5; siphons, 8.0. Females (2 specimens): length, 15.2, 17.1; profemoral length, 4.1, 4.9; siphons, 8.0 (the larger specimen lacks siphons). Color. Female holotype: light brown; lighter mottling not apparent on thoracic or hemelytral dorsa. Second female light brown, lighter mottling evident on dorsum. Male apparently dark brown. Abdominal dorsum brownish red. Abdominal venter dark with four thin, lighter longitudinal bands on sterna 3-6 and a single broad band on parasterna 3-6, except on holotype where parasterna have lighter mottling but no 1996 THE GENUS CURICTA 151 Fig. 17. C. pelleranoi: A) dorsal habitus (5 mm); B) posterolateral metasternal processes, ventral view (0.5 mm); C) male operculum (0.5 mm); D) profemur, anterior view (1 mm); E) parameres (0.25 mm). 152 JOURNAL OF THE NEW YORK ENTOMOLOGICAE SOCIETY Vol. 104(3-4) bands. Protibia with dark basal annulation; no medial annulation. Meso- and meta- thoracic legs concolorous with body. Structural characteristics. Body elongate (Fig. 17A); length 5.0-5.6X maximum width. Eye width 0.5 X interocular distance. Vertex slightly convex; no median carina. Lobe of antennal segment 2, 0.5 X (or slightly more than 0.5 X) length of 3. Pronotum elongate; lateral length approximately 1.2-1.6X posterior width. Trans- verse sulcus incomplete. Median longitudinal sulcus shallow, obsolescent posteriorly. Median longitudinal ridges indistinct, rounded. Lateral sulci shallow. Lateral ridges rounded. Ridges and humeral lobes with black granulations (not apparent on female holotype). Posteroventral extensions of pronotum do not meet in ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width less than length. Trident not apparent; black where handle and transverse parts of lateral prongs would be. Hemelytra widest anterior to base of membranes; black granulations along margins and in clavus. Metathoracic wings fully developed. Last abdominal tergum without carinae or parallel bands of hairs; not tomentose. Prosternum in lateral view not swollen in the posterior half. Mesosternum with midlongitudinal groove present; carinae or bands of hair absent. Metasternum to- mentose; posterolateral processes elongate, slender (Fig. 17B). Parasterna of last abdominal segment ending in small, digitate processes which do not extend poste- riorly beyond the posterior margins of male and female opercula. Male operculum (Fig. 17C). Procoxae 0.5-0.57 X profemoral length. Single, pronounced profemoral tooth on the anteroventral margin of profemoral sulcus; distance from base of profemur to tip of profemoral tooth 0.57-0.60 X total profemoral length. Profemur strongly arched beyond profemoral tooth (Fig. 17D) but not as dramatically as in C. bonaerensis (Fig. 16D). Profemoral sulcus curvilinear, convex posteriorly; no tubercles or sulcal teeth present. Metafemora just reaching anterior margin of abdominal sternum 6 when extended posteriorly. Parameres, Fig. 17E. Female Genitalia: Gp2 arched dorsomedially (Fig. 13D). Discussion. This species is similar to C. bonaerensis. Both species have a single profemoral tooth, a curvilinear profemoral sulcus, tubercles and sulcal teeth absent in the femoral sulcus, and similar male parameres. However, four characters separate these species: in bonaerensis the pronotum is subelongate and the pronotal median sulcus is complete, the prosternum is swollen in the posterior one-half, and the scutellar trident is faint but evident; whereas in pelleranoi, the pronotum is more elongate and the median sulcus obsolescent posteriorly, the prosternum not swollen, and the trident pattern not apparent except for the dark patches noted in the species description above. In addition, all three of the pelleranoi specimens have black gran- ulations on the abdomen and two of them (the female from Sao Paulo and the male) have granulations on the thorax as well. None of the bonaerensis specimens have these markings. Clearly, more collecting of this interesting species will further aid our understanding of the relationship between these two closely related species. Distribution (Fig. 32). Argentina: Corrientes. Brazil: Sao Paulo. Material examined. HOLOTYPE, female: Curicta pelleranoi De Carlo: Argentina, 1996 THE GENUS CURICTA 153 Corrientes, Santo Tome, 10-928. No collector data with specimen but De Carlo indicates in his description (1951:416) that G. Pellerano was the collector. Additional material examined. Two specimens. ARGENTINA. Corrientes: Est. Gal- arza; 4 April 1980; leg. A. O. Bachmann. BRAZIL. Sao Paulo: Cap. Rua Sorocaba 45; 30 March, 1961; leg. C.G.E Curicta peruviana Kuitert Figs. 18, 32 Curicta peruviana Kuitert, 1949:65—66; De Carlo, 1951:419-420; 1960:51. Redescription Measurements. Males: Length, 22.2-23.5; profemoral length, 7. 3-7. 6; siphons, 17.5- 19.0. Females (2 specimens): Length, 25.5-25.8; profemoral length, 8.1; siphons, 19.8-20.3. Color. Uniformly dark brown; no lighter areas evident on thoracic or hemelytral dorsa. Abdominal dorsum brownish red. Abdominal venter dark, no lighter areas evident. Protibia with dark basal annulation; broad, dark medial annulation (one specimen has amber colored protibia which makes it difficult to see the dark medial annulation). Meso- and metathoracic legs uniformly dark. Structural characteristics. Body elongate (Fig. 18 A); length 4. 3-4. 8 X maximum width. Eye width approximately 0.5 X interocular distance. Vertex flat; median, longitu- dinal band of darker hairs that bifurcates anteriorly; posterior crescent-shaped band of darker hairs distinct. Lobe of antennal segment 2 usually >0.5 X length of 3. Pronotum elongate; lateral length approximately 1.3-1.4X posterior width. Trans- verse sulcus nearly complete in some specimens but quite shallow in medial third. Median longitudinal sulcus shallow, obsolescent posteriorly. Median longitudinal ridges prominent, rounded, merging posterior to obsolescent median sulcus, gradu- ally declivent to transverse sulcus. Lateral sulci shallow. Lateral ridges rounded, indistinct. Ridges and humeral lobes covered with small, black granulations. Pos- teroventral extensions of pronotum meet in ventral midline at junction of pro- and mesosterna (Fig. 2B). Scutellum width less than length. Trident not very distinct; vague outline provided by darker hairs. Hemelytra widest Just anterior to base of membrane (females more so than males whose hemelytral margins between humeral lobes and hemelytral membranes tend to be subparallel); dark granulations present, heaviest along margins and in clavus. Metathoracic wings fully developed. Last abdominal tergum of both sexes with distinct pair of parallel, longitudinal bands (one per plate) of dense, long hairs (Fig. 18B). Prosternum in lateral view swollen in the posterior half. Mesosternum with midline groove present; heavily tomentose, appearing granular. Metasternum also heavily tomentose. Parasterna of last abdominal segment ending in broad, ventrally produced processes flanking operculum (Fig. 18C) which posteriorly extend noticeably beyond the posterior margins of the male and female opercula (Fig. 18D). Male operculum (Fig. 18D). 154 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Fig. 18. C. peruviana. A) dorsal habitus (5 mm); B) last abdominal tergum (1 mm); C) last abdominal segment, lateral view (0.5 mm); D) last abdominal segment, ventral view (1 mm), E) profemur, anterior view (2 mm); F) parameres (0.25 mm). 1996 THE GENUS CURICTA 155 Procoxae approximately 0.45 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth approximately 0.44 X total femoral length (Fig. 18E). Profemur moderately arched beyond profemoral teeth. Profemoral sulcus with single row of less than dozen tubercles distally; no sulcal teeth. Metafemora reaching onto posterior half of abdominal sternum 6 when extended posteriorly. Paramere Fig. 18F. Female genitalia: Gp2 dorsoventrally flattened (Fig. 13B). Discussion. Four characters in combination are diagnostic for this species: proximal profemoral teeth; distinct longitudinal bands of long, dense hair on the last abdominal tergum; respiratory siphon length greater than 0.7 X body length; and a broad, dark medial annulus on the protibia. C. pronotata from western Mexico shares the first and last characters with peruviana. However, in pronotata, the last abdominal tergum has a pair of elevated carinae with medial projections (males only; in females this trait is obscure) and the siphons are less than 0.6 X the length of the body. Distribution (Fig. 32). Peru: Sani Beni and Satipo. Material examined. HOLOTYPE, male: Vic. Sani Beni, Peru, 890 m.a.s.l., small pool, 17 Oct. 1935, leg. F. Woytkowski. Deposited in Snow Entomological Museum, University of Kansas. ALLOTYPE, female: same locality data except listed as col- lected in “jungle pools.” Deposited in Snow Entomological Museum, University of Kansas. PARATYPES, four males: same locality data except: three collected in “jun- gle pools”, one in “small pools;” one date given as 19 October 1935. Deposited in Snow Entomological Museum, University of Kansas. Additional material examined. 14 specimens collected in October (12) and November (2); deposited in SEMC. PERU. Sani Beni; Satipo. Curicta hungerfordi Kuitert Figs. 19, 33 (See also 4B, 6) Curicta hungerfordi Kuitert, 1949a:64-65; De Carlo, 1951:396. Redescription Measurements. Males: Length, 12.5-16.8; profemoral length, 3.4-4. 5; siphons, 7.0- 8.2. Females: Length, 14.1-18.9; profemoral length, 4. 1-5.1; siphons, 8. 2-9. 2. Color. Dark brown in larger specimens to light brown in smaller specimens; lighter mottling evident on thoracic and hemelytral dorsa. Abdominal dorsum red. Abdom- inal venter heavily mottled with lighter areas but without longitudinal bands. Protibia dark basally, becoming gradually lighter distally, with or without an indistinct, dark medial annulation. Meso- and metathoracic legs concolorous with body. Structural Characteristics. Body elongate (Fig. 19A): length 3. 8-4. 2 X times maxi- mum width. Eye width about 0.5-0.7X interocular space. Vertex convex; slight median lon- gitudinal Carina evident. Lobe of antennal segment 2 greater than 0.5 X length of 3. Pronotum subquadrate (Fig. 4B); lateral length subequal to posterior width. Trans- verse sulcus complete in most specimens but shallow in medial third of pronotum. Median longitudinal sulcus shallow; usually obsolescent posteriorly, infrequently complete. Median longitudinal ridges prominent, rounded and elevated or subcari- 156 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Fig. 19. C. hungerfordi: A) dorsal habitus (5 mm); B) operculum (0.5 mm); C) profemur, anterior view (2 mm); D) parameres (0.25 mm). 1996 THE GENUS CURICTA 157 nate; fused posterior to obsolescent median sulcus, gradually declivent to transverse sulcus. Paired lateral sulci shallow. Paired lateral ridges rounded, prominent. Ridges and humeral lobes with some granulations; this particularly evident in larger speci- mens. Posteroventral extensions of pronotum meet on ventral midline at junction of pro- and mesosterna (Fig. 2B). Scutellum width subequal to length. Trident pattern not distinct but elevated above rest of scutellum; lateral prongs subcarinate and arched medially; longitudinal parts of lateral prongs and handle usually darker. Hemelytra usually widest just anterior to base of membranes; some of larger spec- imens have subparallel hemelytral margins between humeral lobes and hemelytral membranes; granulations evident along margins and in clavus in larger specimens. Metathoracic wings fully developed. Last abdominal tergum without carinae; parallel, longitudinal bands of longer hairs, one per plate, present in less than one-third of specimens examined. Prosternum in lateral view swollen in the posterior half. Mesosternum with neither midline groove, carinae, nor bands of hair evident. Metasternum tomentose. Para- sterna of last abdominal segment usually ending without terminal processes; a few specimens with short, digitate processes which do not extend posteriorly beyond posterior margins of male and female opercula. Male operculum. Fig. 19B. Procoxae 0.40-0.45 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of antero ventral profemoral tooth 0. 30-0.40 X total profem- oral length (Fig. 19C). Profemur strongly arched beyond profemoral teeth. Profem- oral sulcus with single row of tubercles throughout its length; no sulcal teeth. Metafemora reaching onto proximal third abdominal sternum 6 when extended posteriorly. Parameres, Fig. 19D. Female genitalia: Gp2 arched dorsomedially (Fig. 13D). Discussion. Profemoral teeth occurring at 0.30 to 0.40 X the length of the profemur and the large eye width relative to the interocular distance are diagnostic for this species. Compared to the other two species occurring in Mexico, C. scorpio and C. pronotata, hungerfordi is smaller, has profemoral teeth more proximal, and lacks carinae on the dorsum of the last abdominal tergum. Three characters vary continuously between the smaller and larger specimens in this species. In the smaller specimens (principally a series from Tlalpam, Federal District) the pronotal humeral lobes are not markedly produced posterolaterally (Fig. 4B), eye width begins at approximately 0.5 X the interocular distance, and the dorsum of the body has few apparent granulations. As body size increases, the humeral lobes tend to become more divergent posterolaterally, eye width increases up to approxi- mately 0.7 X the interocular space, and the dorsum of the thorax and abdomen is increasingly likely to have granulations. Finally, it should be noted that two of the three specimens comprising the syntype series for C. scorpio are actually C. hungerfordi (see the discussion for C. scorpio). Distribution (Fig. 33). Mexico: Federal District, Morelos, Jalisco, Michoacan, Hil- dago. Material examined. HOLOTYPE, male: Mexico, Real de Arriba, District of Temas- caltepec, Alt. 1960 m, May-June 1933, leg. H. E. Flinton. Deposited at the Snow Entomological Museum of the University of Kansas. ALLOTYPE, female: Mexico, Real de Arriba, Temascaltepec, 25 May 1933, leg. R. L. Usinger. Deposited at the 158 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Snow Entomological Museum of the University of Kansas. PARATYPES, three fe- males: Mexico, Pachuca, Mor., Hgo Hgw 65 120 km, 21 September 1946, leg. J. G. Shaw (2); Morelos, 14 July 1936. Deposited at the Snow Entomological Museum of the University of Kansas. Additional material examined. 33 additional specimens collected in June (1), July (2), and September (27); deposited in AMNH, ISNB, JTPC, NHRS, SEMC. MEX- ICO. Eederal District: Tlalpam; Xochimilco. Hidalgo: Pachuca, Hwy 65-120 km. Jalisco: 12 km W Ojuelos de Jalisco. Michoacan: Patzcuaro, 7,200', 4 mi. W. Cuidad Hidalgo. Curicta pronotata Kuitert Figs. 20, 33 (See also 3A, 3B, 4A, 5) Curicta howardi [sic], Hungerford, 1922:430-431. (Misidentification) Curicta pronotata Kuitert, 1949a:66-68; De Carlo, 1951:39; Polhemus, J., 1976: 207-208; DuBois, 1978:207; Polhemus, D., 1988:529-530. Redescription Measurements. Males: Length, 19.0-22.5; profemoral length, 5. 8-7.0; siphons, 9.5- 12.7. Females: Length, 20.0-26.0; profemoral length, 6. 5-8. 2; siphons, 10.0-13.8. Color. Grayish-brown; lighter mottling only faintly apparent on thoracic and hem- elytral dorsa. Abdominal dorsum reddish. Abdominal venter with two lighter spots on each of sterna 3-6; no lighter longitudinal bands. Protibia dark basally with wide, dark medial annulation. Distal tibia and tarsus dark. Meso- and metathoracic legs dark. Structural characteristics. Body elongate (Fig. 20A); total length 4. 5-5. OX maxi- mum width. Eye width 0.5 X interocular distance. Vertex convex, median, longitudinal band of darker hairs present, bifurcates anteriorly in about half of specimens; posterior cres- cent-shaped band of darker hairs distinct. Paraclypea noticeably swollen, rounded (Fig. 3 A). Lobe of antennal segment 2 up to 0.5 X length of 3. Pronotum elongate; lateral length 1.1-1.5X posterior width. Transverse sulcus in- complete. Median longitudinal sulcus shallow, obsolescent posteriorly. Median lon- gitudinal ridges very prominent and elevated, quite rounded, fused posterior to ob- solescent median sulcus, gradually declivent to transverse sulcus. Lateral sulci deep. Lateral ridges rounded, prominent. Many small granulations on ridges and humeral lobes; this particularly evident on smaller, less robust specimens. Posteroventral ex- tensions of pronotum meet on ventral midline at junction of pro- and mesosterna (Fig. 2B). Scutellum width less than length. Trident pattern fairly distinct; lateral prongs carinate; middle prong indistinct. Hemelytra with maximum width varying from just posterior to pronotum to ap- proximately midway to base of membranes; small, black granulations throughout, heaviest along membrane margin, claval, and hemelytral lateral margins. Metatho- racic wings short, usually reaching anterior margin of penultimate tergum; infre- quently fully developed. Last abdominal tergum sexually dimorphic: males with 2 parallel, longitudinal 1996 THE GENUS CURICTA 159 Fig. 20. C. pronotata: A) dorsal habitus (5 mm); B) last abdominal tergum (0.5 mm); b) last abdominal segment, lateral view (0.5 mm); c) last abdominal segment, ventral view (1 mm). E) profemur, anterior view (2 mm); F) parameres (0.25 mm). 160 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) carinae (1 per plate) with modest, medially directed swelling, or projection, occurring distally (Fig. 20B); females without carinae, some specimens have two parallel strips of dense hairs, others are simply tomentose on the two tergal plates. Prosternum in lateral view swollen in the posterior half. Mesosternum with midline groove not present; usually tomentose, in some specimens two parallel, longitudinal bands of dense hairs apparent. Metasternum tomentose laterally. Parasterna of last abdominal segment ending in small ventromedial processes flanking operculum (Fig. 20C); posterior margins of male and female opercula usually extend well beyond parasternal processes (Fig. 20D); in dorsal view parasterna project beyond lateral margins of overlying connexiva (Fig. 20A). Male operculum. Fig. 20D. Procoxae 0.42-0.46 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth 0.41 -0.47 X total profe- moral length (Fig. 20E). Profemur moderately arched beyond profemoral teeth. Pro- femoral sulcus with single row of tubercles in distal third to half; no sulcal teeth. Metafemora reaching onto posterior half of abdominal sternum 6 when extended posteriorly. Parameres, Fig. 20F. Female genitalia: Gp2 arched dorsomedially (Fig. 13D). Discussion. The swollen paraclypea, posteromedial swellings on the carinae on the last abdominal tergum in males, and the laterally projecting last abdominal parasterna are diagnostic for this distinctive species. In addition, although a number of species have median pronotal ridges that fuse posterior to an obsolescent median sulcus, these ridges in pronotata are particularly prominent and therefore can also be con- sidered diagnostic. Finally, the occurrence of short metathoracic wings in most spec- imens of this species appears to be unique in the genus. This species is readily separated from the other two species occurring in Mexico, C. hungerfordi and C. Scorpio, by the swollen paraclypea and laterally projecting last abdominal parasterna which are not found in either of these latter two species. Males of scorpio have carinae on the last abdominal tergum but, in contrast to the slender carinae with a posterior, medially directed swelling occurring in pronotata, the carinae of scorpio are more robust and have medially directed swellings occurring midway along the length of the carinae. There is considerable variability in pronotal (Fig. 4A), antennal (Fig. 3B), and paramere (Fig. 20F) shape in this species. Forms with a more robust pronotum generally have shorter respiratory siphons and more elongate lobes on antennal seg- ment 2 and are found in southern Arizona and northern Sonora, while more slender specimens with longer respiratory siphons and shorter antennal lobes tend to be found at the southern extreme of the species’ range at La Quemada, Nayarit. How- ever, there does not appear to be clinal variation in these characters, as the specimens occurring at other localities do not exhibit either incremental or continuous change with latitude. For example, the least robust specimen was collected at the northern- most part of the species range outside Tucson, Arizona; and specimens collected around Tepic, Nayarit, have widely divergent, robust humeral lobes but are quite narrow more anteriorly. Specimens collected in southern Sonora localities exhibit no particular tendency in pronotal robustness, with some specimens resembling the northern robust form and others the more slender La Quemada specimens. Clasper variation exhibits no trend at all. In sum, it would appear that this is a particularly variable species, but that variability exhibits no obvious geographic pattern. Further 1996 THE GENUS CURICTA 161 collecting of this distinctive species throughout its range, with particular emphasis on Sinaloa where only one specimen has been taken, will further elucidate the pattern of variability. Distribution (Fig. 33). United States: southern Arizona. Mexico: Sonora, Chihuahua, Sinaloa (?), Nayarit. Material examined. HOLOTYPE, male: U.S.A.: Huachuca Mts. No collector or tem- poral data. Deposited at the Snow Entomological Museum of the University of Kan- sas. PARATYPES, two males: U.S.A: Huachuca Mts. No collector data, one speci- men dated 1899. Deposited at the Snow Entomological Museum of the University of Kansas. One of the paratypes has a “C. howardi, det. Drake” label on it. Kuitert notes in his original description of this pronotata that Hungerford relied on this misidentification by Drake for his 1922 redescription of ""howardi.'' Additional material examined. 70 specimens collected in April (6), May (6), June (9), July (1), August (25), September (1), October (1), November (4), and December (17); deposited in ASU, CINCB, JTPC, LACM, and UA. U.S.A. Arizona: Sabino Canyon (nr. Tucson); Ramsey Canyon, 1820m, Huachuca Mtns., Cochise Co. MEX- ICO. Sonora; Canyon de Evans, Sierra de los Ajos; El Cobre; 1 mi. E. Maicova; 10 and 11 mi. E. Yecora; 17 mi. S. Bacanora. Chihuahua: Sierra Madres, 3 mi. W. Rio Negro. Sinaloa: Sierra Suratato, 7300', La Bufa Suratato (No such locality has been located. John Polhemus believes the site is probably in southwestern Sonora, pers. comm.). Nayarit: 20, 22, 23, 26, and 30 mi. SE Tepic. Jalisco: La Quemada. Curicta scorpio Stal Figs. 21, 22, 34 (See also, 2B, 2D, 9B, 11, 12, 13D) Curicta scorpio StM, 1862 (1861):203 [Lectotype here designated]; Stal, 1865:185; Champion, 1901:352-353; Montandon, 1903:98; Kirkaldy and Torre Bueno, 1909: 202; De Carlo, 1951:395; Keffer, 1993:434-435. Nepa scorpio, Ferrari, E. von, 1888:191 [Champion, 1901:353, placed scorpio back in Curicta]. Nepoidea montandoni Martin, 1898:68 [Syn. by Champion, 1901:353]. Curicta howardi Montandon, 1909b (1910):181-183 [Syn. by Keffer, 1993:434— 435]; Kuitert, 1949a:68; De Carlo, 1951:395; Polhemus, J., 1976:208; Polhemus, D., 1988:529. Curicta drakei Hungerford, 1922:432-433 [Syn. by Kuitert, 1949a:68]; De Carlo, 1951:395; Gonsoulin, 1975:25. Redescription Measurements. Males: Length, 16.0-20.5; profemoral length, 4. 5-5. 5; siphons, 6.2- 9.0. Females: Length, 18.4-22.8; profemoral length, 4. 8-5. 7; siphons, 6. 7-9. 5. Color. Dark brown; lighter mottling on thoracic and hemelytral dorsa evident. Ab- dominal dorsum red. Abdominal venter heavily mottled with lighter areas but without longitudinal bands. Protibia dark basally; medially variable, with or without dark medial annulus; when present irregular, i.e., may lighten distally, be indistinct, or even appear as two indistinct annuli. Meso- and metathoracic legs dark. Structural characteristics. Body elongate (Fig. 21 A); length 4. 5-5. 5 X maximum width. 162 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Eig. 21. C Scorpio: A) dorsal habitus (5 mm); B) last abdominal tergum (0.5 mm); C) last abdominal segment, lateral view (0.5 mm); D) last abdominal segment, ventral view (0.5 mm); E) male operculum (0.5 mm); E) profemur, anterior view (2 mm); G) parameres (0.25 mm). 1996 THE GENUS CURICTA 163 22 A B Eig. 22. C. Scorpio: A) head and prothorax (Oaxaca, Mexico), lateral view (1 mm); B) head and prothorax (Texas, U.S.), lateral view (1 mm). Eye width approximately 0.5 X interocular distance. Vertex convex; prominent median longitudinal carina (Figs. 22A, B) with anterior bifurcation apparent infre- quently; crescent-shaped band of hair posteriorly also evident. Lobe of antennal segment 2 usually greater than 0.5 X length of 3. Pronotum elongate; lateral length approximately 1.1-1.3X posterior width. Trans- verse sulcus incomplete. Median longitudinal sulcus deep to moderately deep and complete, in some specimens becoming shallower posteriorly. Median longitudinal ridges prominent, carinate. Lateral sulci deep. Lateral ridges rounded. Entire pro- notum heavily tomentose, appearing very granular on ridges and humeral lobes. Posteroventral extensions of pronotum meet on ventral midline at junction of pro- and mesosterna (Fig. 2B). Scutellum width less than length. Trident distinct; carinate and usually darker than rest of scutellum. Hemelytra widest just anterior to base of membranes (females more so than males whose hemelytral margins between humeral lobes and hemelytral membranes may be subparallel); moderately to heavily covered with granulations particularly along lateral margins and in clavus. Metathoracic wings fully developed. Last abdominal tergum sexually dimorphic: males with two robust, parallel, lon- gitudinal carinae (one per plate) with a prominent medial swelling, or projection, occurring midlength (Fig. 2 IB); females with straight carinae without medial pro- jection, less robust. Prosternum in lateral view variable: swollen in posterior half (Fig. 22A) to decli- vent from anterior to posterior (Figs. 22B). Mesosternum with midline groove prom- inent; tomentose; two parallel, longitudinal ridges, slightly elevated occurring later- 164 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) ally. Metasternum heavily tomentose. Parasterna of last abdominal segment without apparent distal processes (Figs. 2 1C, D); posterior parasternal margins extend sub- equally to posterior margins of male and female opercula. Male operculum, Fig. 21E. Procoxae 0.40-0.46 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth 0.40-0.45 X total profe- moral length (Fig. 2 IF). Profemur moderately arched beyond profemoral teeth. Pro- femoral sulcus with a single row of tubercles usually throughout length of sulcus; no sulcal teeth. Metafemora reaching onto proximal fourth of abdominal sternum 6 when extended posteriorly. Parameres Fig. 21G. Female genitalia: Gp2 arched dorsomedially (Fig. 13D). Discussion. Prominent carina on the vertex, proximal profemoral teeth, and two parallel carinae on the last abdominal tergum are, in combination, diagnostic for this species. C. scorpio shares with pronotata proximal profemoral teeth and carinae on the last abdominal tergum. However, in scorpio the carinae are prominent in both sexes and, in the male, there is a medially directed swelling midway along the length of each carinae, whereas in pronotata the carinae are very faint in females and the swelling in males occurs distally. In addition, scorpio is heavily tomentose and cov- ered with granulations on its dorsum and thoracic sternum; pronotata is less tomen- tose and has fewer, and smaller granulations. Finally, scorpio lacks pronotata's swol- len paraclypea. C. scorpio can be differentiated from the other species occurring in Mexico, C. hungerfordi, by the above mentioned abdominal carinae, which are ab- sent in hungerfordi. Finally, it is worth noting that the synonymy of C. drakei by Louis Kuitert was rather informal. Writing at the end of the discussion of his new species, C. pronotata, he stated that one of his pronotata type specimens had previously been incorrectly determined by Carl Drake as C. howardi. Thus, says Kuitert, Hungerford, relying on this determination, then went on to describe drakei, actually howardi, as a new species. Distribution (Fig. 34). United States: southern Louisiana, southeastern Texas. Mex- ico: Tamaulipas, Vera Cruz, Nayarit, Jalisco, Colima Michoacan, Guerrero, Morelos, Federal District, Oaxaca, Chiapas, Mexico. Guatemala. Honduras. Nicaragua. This is the only curictan species with specimens found both in North and Central America. All other species have either a South American or a North American distribution. Material examined. LECTOTYPE, female: Curicta scorpio Stal. Mexico, Salle. De- posited in the Rijksmuseum Van Natuurlijke Historie, Stockholm. This specimen bears a red type label. Two other specimens were loaned from Stockholm with this holotype and the three specimens together were called syntypes on the museum loan invoice. All three specimens bear two small typed or printed labels saying, “Mexi- co” and “Salle.” The scorpio type also has a printed label which reads “sp. Eig- ured.” referring, presumably, to the Eigure 1 in Champion, 1901 (Stal did not provide figures with his original description). There is no way of knowing how many spec- imens StM looked at, his original description being mute on this matter, although he gives only one set of measurements. What makes all this so interesting is that the other two specimens are clearly hungerfordi and not scorpio. Both these hungerfordi specimens have printed labels indicating that H. B. Hungerford looked at them during 1996 THE GENUS CURICTA 165 his 1928 European trip and one of the specimens has a Hungerford handwritten determination label reading, “C. scorpioT’’ Apparently, Hungerford had his doubts as to whether these three specimens were a good series. It remained for his student, Louis Kuitert, studying specimens collected subsequent to Hungerford’s European trip and deposited in SEMC, to describe hungerfordi, which matches these two Stockholm specimens. Kuitert does not mention the Stockholm specimens in his description of C. hungerfordi. In order to avoid confusion in the future over these specimens, I will assume that the three specimens did originally constitute a series as the curatorial staff in Stockholm believes. Therefore, I hereby designate the Scor- pio specimen as a lectotype. The other two specimens now bear my determination label as C. hungerfordi. HOLOTYPE, male: Nepoidea montandoni Martin. Mexique. Salle. Bears a handwritten determination label reading, “Nepoidea, Montandoni, type J. Martin, 23 fev. 1898.” There is also a printed label reading “type.” Deposited in the Museum National D’Histoire Naturelle, Paris. I do not know if this Salle spec- imen might originally have been part of the Stockholm Salle series discussed above. It is clear from Martin’s original description that he looked at one specimen so I am affixing a holotype label to the specimen. HOLOTYPE, female: Curicta howardi Montandon. Victoria, Texas. 12 June 1904. There is no collector label. Montandon dedicates the naming of the specimen to L. O. Howard but there is no indication that Howard was the collector. The case for synonymizing this species with scorpio is discussed in detail in Keffer, 1993. There is a handwritten determination label reading, “Curicta, Howardi, Montandon, type 1909 Dec.” Deposited in the National Museum of Natural History, Washington, D.C., type no. 26224. HOLOTYPE, male: Curicta drakei Hungerford. Rock Island Co., Texas. 30 August 1922, leg. Grace Wiley. Deposited in the Snow Entomological Museum at the University of Kansas. ALLOTYPE, female: Curicta drakei Hungerford. Same locality, temporal, and col- lector data as holotype. Deposited in the Snow Entomological Museum at the Uni- versity of Kansas. PARATYPES, 10 females; five males: Curicta drakei Hungerford: Colorado Co., Texas, 22 August 1922 (8), leg. Grace Wiley; Colorado Co., Texas, 24 June 1922, leg. Grace Wiley (1); Colima, Mexico, leg. L. Conradt (no temporal data) (1); New Orleans, La., May, 1916, no collector data (2). Deposited at the Snow Entomological Museum of the University of Kansas (12) and in the general type collection (1) and in the Carl Drake Collection (2), both of the National Museum of Natural History, D.C. One of the Kansas paratypes bears a R. I. Sailer determination label identifying the specimen as C. howardi, “compared with type.” The drakei paratype in the general type collection of the National Museum of Natural History, Washington, D.C. bears a similar determination label by Sailer. Additional material examined. 310 specimens collected in the United States in March (1), April (3), June (3), July (58), August (179), September (3), October (6), and in Mexico to Nicaragua in February (1), March (2), May (2), June (13), July (15), August (16), October (1), and December (7); deposited in USNM, JTPC, SEMC, CAS, AMNH, LSU, LACM, TAM, SIU, USU, UCD, and ERIAC. U.S.A. Texas: Colorado Co., Skull Creek; Galveston Co., nr. Hitchcock; Brazos Co., 8 mi W Bryan; College Station; Postoak Lake; Little Brazos River; Live Oak Co., 10 mi. SWGeorge West; Liberty Co., 4 mi. W Moss Hill; Jim Wells Co.; Travis Co., Austin; Gonzales Co., Ottine Palmetto St. Pk; Jackson Co., Ganado; Walker Co., Sam Houston State Park. Louisiana: Iberville Par., Gabriel Exp Sta.; Roux Lafourche Par.; Charles Par., 166 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) trib. Lanaux Canal at US 90E jet. Hwy 3060; Baton Rouge; New Orleans. MEXICO. Tamaulipas: 5 mi S Ciudad Victoria; 1/4 mi N San Jose de las Rusas. Vera Cruz: 15 mi W and 20 mi S Vera Cruz; near Perote; 15 mi NW Acayucan; 3 mi W Paso de Ovejas; 13 km WNW Potrero. Nayarit: Tepic; Compostela; N of Compostela. Michoacan: El Sabino, Uruap.; Patzeuaro. Jalisco: Atenquique. Colima: Colima. Guerrero: Rio Balsas jet Ocapulca. Morelos: 5 mi E Cuernavaca; Temixco; Rio Amacuza, 133 km S Mexico City. Oaxaca: Tapanatepec, 17 mi E Oax. Chiapas: Ocozocoautla; W. Rizode de Oro. San Luis Potosi: C. Valles. Villa Warez Valles (state unknown). GUATEMALA: E of Quirigua. HONDURAS: Escuela Agricola Panamericana; 1 km N Las Moras; 2 km S Las Capucas Que Brada de Bijao. NIC- ARAGUA: Managua; 6 mi N Managua; 22 mi S Rivas. Curicta grandis De Carlo Figs. 23, 35 Curicta grandis De Carlo, 1951:396—397. Curicta paraguayensis De Carlo, 1951:397-399. NEW SYNONYMY. Curicta borellii [sic], De Carlo, 1951:399-401. (Misidentification) Redescription Measurements. Males: Length, 26.3-29.0; profemoral length, 7. 1-8.3; siphons, 21.0- 27.0. Females: Length, 29.2-32.5; profemoral length, 7. 5-8. 5; siphons, 23.0-31.0. Color. Dark brown; lighter mottling evident on the thoracic and hemelytral dorsa. Abdominal dorsum reddish brown. Abdominal venter with lighter mottling, infre- quently aggregating into two longitudinal bands on sterna 3-6. Protibia dark basally; no medial annulation; protibia is dark at the junction with the protarsus which is also dark. Meso- and metathoracic legs golden with some dark mottling. Structural characteristics. Body elongate (Fig. 23 A); length 5. 0-5. 3 X maximum width. Eyes large; width 0.5-0.6X interocular distance. Vertex slightly convex; ru- dimentary median, longitudinal band of darker hairs present posteriorly, rarely con- tinues anteriorly but when it does anterior bifurcation present; crescent-shaped band of hairs evident posteriorly. Lobe of antennal segment 2 elongate, usually just a little shorter than length of 3. Pronotum elongate; lateral length 1.3-1. 4X posterior width. Transverse sulcus in- complete. Median longitudinal sulcus very shallow, sometimes complete, usually obsolescent posteriorly. Median longitudinal ridges prominent, elevated, rounded. Lateral sulci very shallow. Lateral ridges rounded, not pronounced. A few black granulations on ridges and humeral lobes, but these not pronounced. Posteroventral extensions of pronotum meet on ventral midline at junction of pro- and mesosterna (Fig. 2B). Scutellum width less than length. Trident pattern distinct; area delimited by lateral prongs elevated above rest of scutellum; all parts of trident carinate and darker. Hemelytra widest just posterior to humeral lobes; granulations along elytral lateral margins. Metathoracic wings fully developed. Last abdominal tergum tomentose usually with two distinct subparallel mediolon- gitudinal bands of long hairs (1 per plate). Prosternum in lateral view dramatically swollen in posterior half (Fig. 23B). Me- 1996 THE GENUS CURICTA 167 Fig. 23. C. grandis: A) dorsal habitus (5 mm); B) head and prothorax, lateral view (3 mm); C) last abdominal segment lateral view (0.5 mm); D) last abdominal segment, ventral view (1 mm); E) male operculum (0.5 mm); F) profemur, anterior view (2 mm); G) parameres (0.25 mm). 168 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) sosternum with midline groove and two broad, parallel, longitudinal bands of dense, short hair present. Metathoracic sternum tomentose. Parasterna of last abdominal segment ending in very broad processes (Fig. 23C), ventrally produced, which pos- teriorly extend noticeably beyond the posterior margins of the male and female opercula (Fig. 23D). Male operculum. Fig. 23E. Procoxae slightly less than 0.50 X profemoral length. Two profemoral teeth; dis- tance from base of profemur to tip of anteroventral profemoral tooth just a little over 0.50 X total profemoral length (Fig. 23F). Profemur only slightly arched beyond profemoral teeth. Profemoral sulcus with incomplete row of tubercles on distal sixth; no sulcal teeth distally. Metafemora reaching onto anterior half abdominal sternum 6 when extended pos- teriorly. Parameres, Fig. 23G. Female genitalia; Gp2 dorsoventrally flattened (Fig. 13B). Discussion. The dramatically swollen prosternum, distal profemoral teeth, absence of profemoral sulcal teeth, and the large size of the body and siphons are diagnostic for this species. Only peruviana approaches the size of grandis, has a swollen pro- sternum, and also lacks sulcal teeth in the profemora. However, peruviana is usually smaller in both sexes, is less dramatically swollen in the prosternum, has the pro- femoral teeth more proximal (as opposed to halfway in grandis), and has a broad medial annulus on the protibia which is lacking in grandis. The history of this species is closely linked with that of borellii. De Carlo de- scribed two species with the same general facies as his understanding of ''borellii," C. grandis and paraguayensis. C. grandis was said to resemble "borellii" in pron- otal, prosternal, and mesosternal characteristics, and to differ from "borellii" by being larger with longer siphons (De Carlo, 1951:395). Yet none of the "borellii" specimens in De Carlo’s collection match Montandon’s original (1903) description for borellii. Instead, these specimens, and also De Carlo’s type for paraguayensis, do match the description given here for grandis. They dis- play the characteristic swollen prosternum of grandis while lacking the profemoral sulcal teeth of borellii. Nevertheless, except for the types of grandis and paraguay- ensis, De Carlo (1951) believed all his specimens to be Montandon’s borellii. In contrast, the single available specimen that was determined by Montandon to be borellii (the syntypes for borellii have not been located) does indeed match Mon- tandon’s original (1903) borellii description. De Carlo’s "borellii" specimens do, however, resemble De Carlo’s own type for grandis (and for paraguayensis, see below), at least within the range of variability found intraspecifically in other species of Curicta. Thus, the swollen prosternum as an attribute of "borellii," reported by De Carlo and repeated by Kuitert (1947: 43-44, unpub.), appears to be erroneous. De Carlo’s "borellii" specimens, which have a swollen prosternum and lack pro- femoral sulcal teeth, are therefore here determined to be grandis. For a more detailed account of this confusion, see the discussion of C. borellii. According to De Carlo (1951), C. paraguayensis differed from grandis because of a narrower prothorax, less convex pronotum, more distinct longitudinal sulci on the pronotum, more slender profemora, and a slightly wider body (ibid: 399). Both species, grandis and paraguayensis, were also said to differ from each other (as well as from "borellii") in antennal shape. However, all the structural differences cited by De Carlo between grandis and paraguayensis are slight and well within the range 1996 THE GENUS CURICTA 169 of variability found intraspecifically in other species of Curicta. I therefore synon- ymize paraguayensis with grandis. In the absence of priority for either name, I elect C grandis for this species because of its robust form. Distribution (Fig. 35). Northern Argentina, Paraguay, Bolivia, and Suriname. This species has a curiously disjunct distribution with the southern and northern popula- tions separated by the wide expanse of the Amazon Basin. Further collecting is needed to determine if C. grandis is truly absent from the Amazon. Material examined. HOLOTYPE, female: C. grandis De Carlo. Girardet, S. del Es- tero, Argentina. Collected in June, day and year illegible. No collector label but De Carlo obtained the specimen from Juan M. Bosq (ibid: 397). HOLOTYPE, female: C. paraguayensis De Carlo. Villarrica, Paraguay. April, 1938. No collector label but De Carlo stated that the specimen was obtained from Federico Schade (ibid: 398). Additional material examined. 96 specimens collected in February (2), March (18), May (28), June (11), July (17), August (1), September (9), November (1), and De- cember (2); deposited in JTPC, MACN, MZSP, NMNH, and SEMC. ARGENTINA. Salta: Tartagal; Acambuco. Formosa: Ing. Juarez. Santiago del Estero: Las Juries. Chaco: Sante Fe. Sante Fe: Fires Lille. PARAGUAY. Concepcion: Vallemi. Guaira: Villarrica. Chaco: P. N. Defensores del Chaco, Madrejon.: Estancia Postilion Puerto Max at Rio Paraguay (province unknown). BOLIVIA. Santa Cruz: Santa Cruz. SU- RINAME: Marowijne: Christian Kondre. Curicta tibialis (Martin) Figs. 24, 35 Nepoidea tibialis Martin, 1898:66. Curicta tibialis, Montandon, 1903:98; Kirkaldy and Torre Bueno, 1909:202; Mon- tandon, 1909a:144; De Carlo, 1951:401-402. Curicta suspecta Montandon, 1903:99-100. Syn. by Montandon, 1909a: 144. Curicta beckeri De Carlo, 1956:6. NEW SYNONYMY. Redescription Measurements. Males: Length, 26.8-28.0; profemoral length, 6. 7-7. 4; siphons, 19.5 {beckeri HT with broken siphons-not measured). Females: Length, 29.3-31.3; pro- femoral length, 7.5-8. 1; siphons, 20.1-24.8. Color. Grayish brown. Lighter mottling on thoracic and hemelytral dorsa evident. Abdominal dorsum dark reddish brown. Abdominal venter with lighter, irregular longitudinal bands on sterna and parasterna 3-6. Protibia dark basally; medial an- nulation variable; complete (two specimens), partial (one), absent (five). Meso- and metathoracic legs dark. Structural characteristics. Body elongate (Fig. 24A); length 5. 2-5. 5 X maximum width. Eye width approximately 0.5 X interocular distance. Vertex convex, with slight to pronounced median longitudinal carina. Lobe of antennal segment 2 usually slightly less than length of 3. Pronotum elongate; lateral length approximately 1.3-1. 4X posterior width. Trans- verse sulcus incomplete. Median longitudinal sulcus deep, becoming shallow or ob- solescent posteriorly. Median longitudinal ridges prominent, rounded. Lateral sulci 170 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Eig. 24. C. tibialis: A) dorsal habitus (5 mm); B) operculum (0.5 mm); C) profemur, anterior view (2 mm); D) parameres (0.25 mm). 1996 THE GENUS CURICTA 171 deep. Lateral ridges rounded, prominent. Few granulations evident on ridges and humeral lobes. Posteroventral extensions of pronotum do not meet on ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width less than length. Trident pattern distinct, all parts carinate and darker. Hemelytra with lateral margins subparallel from pronotum to base of membranes; granulations faintly apparent along lateral margins. Metathoracic wings fully devel- oped. Last abdominal tergum tomentose with longer hairs medially forming indistinct mediolongitudinal bands. Prosternum in lateral view not swollen in the posterior half. Mesosternum tomen- tose; midline groove present in some specimens and apparently absent in others; broad, hairy, rather indistinct, parallel bands present in one specimen, absent in others. Metasternum tomentose. Parasterna of last abdominal segment ending in large, thumb-like processes (as in Fig. 29B) which, in all but one specimen, extend posteriorly slightly beyond posterior margins of male and female opercula. Male operculum. Fig. 24B. Procoxae approximately 0.50X profemoral length. Two profemoral teeth; distance from base of profemur to tip of antero ventral profemoral tooth 0. 5 0-0. 60 X total femoral length. Profemur moderately arched beyond profemoral teeth. Profemoral sulcus, in lateral view, notched on distal sixth, appearing to “step-down” (Fig. 24C); complete row of tubercles; two prominent sulcal teeth distally; tubercles form a single row proximally but proliferate on the two sulcal teeth. Metafemora reaching onto anterior half of abdominal sternum 6 when extended posteriorly. Parameres, Fig. 24D. Female genitalia: Gp2 dorsoventrally flattened (Fig. 13B). Discussion. The large body size and distal notch in the profemoral sulcus together are diagnostic for this species. Only one other species, C. grandis, attains a com- parable body length. However, C. grandis lacks profemoral sulcal teeth and the step- down feature of tibialis' profemoral sulcus. De Carlo thought his beckeri to be the same as tibialis in body and siphon lengths, and prosternal and profemoral attributes. He differentiated these two species only on the basis of differences in antennal and pronotal shape, and the vertex. The differ- ences between the types in the two former characters are small and well within the limits of variability found for other species described in this review of Curicta. The reference to vertex differences is puzzling. According to De Carlo, tibialis has a midlongitudinal carina on the vertex but his species, beckeri, does not. This character is difficult to evaluate on the holotype for beckeri because that specimen has been abraded or chemically treated on the head, pronotum, and scutellum, leaving a hair- less and narrowly convex vertex. All the other specimens of the species have a carina on the vertex, accentuated by a band of upright hairs, which vary in their promi- nence. Interestingly, the specimen with the least prominent carina is a female iden- tified by De Carlo as tibialis. Another female, identified by De Carlo as beckeri, has a very noticeable carina. In sum, the differences between the specimens for this trait are not significant and I therefore am confident in synonymizing beckeri with tibialis. Distribution (Fig. 35). Brazil: Rio Grand do Sul. Uruguay: Durazno. Paraguay(?): Concepcion. Material examined. HOLOTYPE, female: Nepoidea tibialis Martin. Locality: Rio 172 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Grande; Coll. G. Fallon, 259-95; no temporal data or information as to who actually collected the specimen. Deposited in the Museum National D’Histoire Naturelle, Paris. The locality given for this specimen is simply, “Rio Grande,” which probably referred to Rio Grande do Sul, Brazil. This specimen bears a handwritten determi- nation label with the species name, the word “type,” the name “J. Martin,” and the date “28 fe’r 1898.” It is clear from Martin’s original description that he had a single specimen (“un exemplaire de la collection G. Fallou”, p. 66) therefore the specimen I have before me must be the holotype, and I have so labeled it. HOLO- TYPE, female: Curicta suspecta Montandon. Locality: S. Leopoldo. No collector label or temporal data. Determination label by Montandon, dated 1903. Deposited in the Naturhistoriska Riksmuseet Stockholm (NRS). S. Leopoldo probably refers to Sao Leopoldo in Rio Grande do Sul, Brazil. It is clear that Montandon studied only a single specimen when first describing suspecta (“un exemplaire unique”, p. 99) and therefore the specimen I have before me from the Stockholm Museum with Montandon’s determination label and a “typus” label must also be a holotype and I have so labeled it. HOLOTYPE, male: Curicta heckeri De Carlo. Brazil, Rio Gran- de do Sul, Osorio; May (5), 1950; leg. J. Baker. De Carlo lists the collection date for this specimen as “1-1950” (p. 6) but the handwritten label indicates a number 5 for the month. In his original description, De Carlo states that a microscope prepa- ration #3273 exists (this probably was of the genital parameres since none are to be found on the specimen’s phallus). I have received 1 1 microscope slides from the MACN, De Carlo’s home museum, but none by that number. Presumably the slide exists somewhere in the MACN. Additional material examined, four specimens collected in April (1) and July (1) (two specimens without temporal data); deposited in HNHM, MACN, and RMNH. BRAZIL. Rio Grande do Sul: Sao Leopoldo. URUGUAY. Blanquillo, Durazno. PAR- AGUAY(?). Concepcion (this locality is given without a country). Curicta carinata Kuitert Figs. 25, 35 Curicta carinata Kuitert, 1949a:62-64; De Carlo, 1956:6; 1960:51. Curicta dilatata De Carlo, 1951:412-414. [Syn. by De Carlo, 1960:51.] Redescription Measurements. Males: Length, 17.5-20.0; profemoral length, 5. 2-5. 8; siphons, 10.0- 12.2. Females: Length, 17.3-24.0; profemoral length, 5. 5-7.0; siphons, 11.5-15.0. Color. Dark brown. Lighter mottling evident only on the humeral lobes of the pro- thorax. Abdominal dorsum brownish red. Abdominal venter lighter brown with light- er mottling and faint indications of longitudinal bands on sterna and parasterna 3- 6. Protibia with dark basal annulation; distinct, dark medial annulation. Meso- and metathoracic legs golden. Structural characteristics. Body elongate (Fig. 25 A); length approximately 4.5— 5.7 X maximum width. Eye width 0.5 X interocular space. Vertex with prominent medial, longitudinal Carina (Fig. 25B). Lobe of antennal segment 2 usually greater than 0.5 X length 3. Pronotum elongate; lateral length approximately L4-1.5X posterior width. Trans- 1996 THE GENUS CURICTA 173 Fig. 25. C. carinata: A) dorsal habitus (5 mm); B) head and anterior prothorax, lateral view (1 mm); C) meso- and metasterna (1 mm); D) last abdominal segment, lateral view (0.5 mm); E) last abdominal segment, ventral view (0.5 mm); F) profemur, anterior view (2 mm); c) parameres (0.25 mm). 174 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) verse sulcus incomplete. Median longitudinal sulcus deep to moderately deep, com- plete. Median longitudinal ridges prominent, carinate. Lateral sulci deep. Lateral ridges subcarinate. Ridges and humeral lobes densely covered with granulations. Posteroventral extensions of pronotum do not meet on ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width less than length. Trident pattern distinct; longitudinal section of lateral prongs, medial prong, and handle carinate and black; transverse section of lateral prongs lighter, indistinct. Hemelytra widest just anterior to elytral membranes (more so in females than males); large, dark granulations on entire hemelytral surface, heaviest along lateral margins and in clavus. Metathoracic wings fully developed. Last abdominal tergum tomentose with pair of parallel, mediolongitudinal bands of long hairs. Prosternum in lateral view not swollen in posterior half. Mesosternum without midline groove; two prominently elevated, narrow, parallel, longitudinal carinae present (Fig. 25C). Metasternum subglabrous medially, tomentose around lateral and posterolateral margins. Parasterna of last abdominal segment ending in slender dig- itate processes (tending to be larger, less slender in females) (Fig. 25D), which extend subequal to slightly beyond the posterior margins of male and female opercula (Fig. 25E). Male operculum. Fig. 25E. Procoxae about 0.45-0.55 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth approximately 0.47- 0.52 X total femoral length (Fig. 25F). Profemur moderately arched beyond profe- moral teeth. Profemoral sulcus with a single row of tubercles in distal third which are often obscured by the bands of short, dense hair flanking sulcus; no sulcal teeth. Metafemora reaching approximately halfway onto abdominal sternum 6 when ex- tended posteriorly. Parameres, Fig. 25G. Female genitalia: Gp2 dorsoventrally flattened (Fig. 13B). Discussion. This a very distinctive, and homogeneous, species. The pronounced ca- rinae on the mesosternum and vertex, dark medial protibial annuli, heavily granular dorsum, and the absence of sulcal teeth on the profemora are, in combination, di- agnostic. C. cahnata might conceivably be confused with two other species, gran- ulosa and decarloi, since all three species have carinae on the vertex and protibial annuli. C. granulosa also has a heavily granular dorsum but, unlike carinata, has profemoral sulcal teeth and broad tomentose parallel bands on the mesosternum rather than the distinctly elevated carinae of carinata. The respiratory siphons of decarloi are approximately 0.9 X the body length, whereas the siphons of carinata attain a maximum of 0.7 X body length. Both decarloi and carinata lack profemoral sulcal teeth but the tubercles in the profemoral sulcus distally of decarloi are irregular while in carinata the row of tubercles is regular throughout. Also, decarloi has slight carinae on the mesosternum and not the pronounced carinae of carinata. Finally, decarloi apparently lacks the heavy granulation on the thoracic and hemelytral dorsa found in carinata. Distribution (Fig. 35). Paraguay, Bolivia, Peru, Ecuador, Colombia, Trinidad, and Panama. This is the only curictan species with specimens found both in South and Central America. All other species have either a South American or a North Amer- ican distribution. 1996 THE GENUS CURICTA 175 Material examined. HOLOTYPE, male: Curicta carinata Kuitert: Caravene, Para- guay; 15 February 1925; leg. F. Schade. Deposited in the Snow Entomological Mu- seum, University of Kansas. ALLOTYPE, female: Curicta carinata Kuitert: Villar- rica, Paraguay [separate label reads, Coraveni]; 21 September 1924; leg. Fran. Schade. Deposited in the Snow Entomological Museum, University of Kansas. PARATYPES, three males, three females: Curicta carinata Kuitert: 4 collected in Villarrica, Paraguay [Coraveni] on 10 April 1923, 18 April 1923, 11 September 1923, 12 January 1924; 1 collected in Caraveni, 15 June 1924; and 1 collected in Villarico, 19 January 1926. I assume the differences in spellings for Coraveni and Villarrica are typographical errors and do not reflect different localities. Deposited in the Snow Entomological Museum, University of Kansas HOLOTYPE, female: Curicta dilatata De Carlo: Villarrica, Paraguay; 19 May 1938; leg. E Schade. De- posited in Museo Argentino de Ciencias Naturales “Bernadino Rivadavia” ADDI- TIONAL Material Examined. 140 specimens collected in January (75), February (3), March (5), April (11), May (1), June (15), August (3), September (16), November (2), and December (1); deposited in AMNH, ESC A, JTPC, NMNH, NNC, SEMC, and ZMH. PARAGUAY. Villarrica: Villarrica, Coraveni. Aregua, 20 mi E Asuncion. BOLIVIA. Santa Cruz: Neuva Moka. El Beni: 2 km E San Borja. La Paz: 3 km SE Sapecho. PERU. Madre de Dios: Parque Mana, Pakitza. ECUADOR. Napo-Pastaza: Santa Cecilia, 340 m. COLOMBIA. Antioquia: Chigorodo, Rio Leon. TRINIDAD: Chatham. PANAMA. Panama: 3 km SE Ipeti; 3 km S of Ipeti. Herrara: N. fork of Rio Paritas. Description Curicta decarloi, new species Figs. 26, 35 Measurements. Female (1): Length, 25.5; profemoral length, 7.3; siphons, 22.0 Color. Largely obscured by dirt, appears dark reddish brown; lighter markings ob- scured on both dorsum and venter. Abdominal dorsum reddish brown. Legs concol- orous with body. Structural characteristics. Body elongate (Fig. 26 A); total length 5.4X maximum width. Eye width 0.5 X interocular distance. Vertex convex with a pronounced midlon- gitudinal Carina. Lobe of antennal segment 2 slightly greater than 0.5 X length of 3. Pronotum elongate; lateral length 1.5X posterior width. Transverse sulcus incom- plete. Median longitudinal sulcus complete, uniformly deep. Median longitudinal ridges carinate. Lateral sulci deep. Lateral ridges carinate. Granulations not apparent on ridges or humeral lobes. Posteroventral extensions of pronotum do not meet on ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width less than length. Trident distinct; carinate; longitudinal section of lateral prongs, medial prong, and handle black, transverse section of lateral prongs not dark, indistinct. Hemelytra widest just anterior to base of membrane; without apparent granula- tions. Metathoracic wings fully developed. Last abdominal segment tomentose, paired carinae or bands of hair not apparent. Prosternum in lateral view not swollen. Mesosternum without midline groove; 176 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) D Eig. 26. C. decarloi: A) dorsal habitus (5 mm); B) last abdominal segment, lateral view (0.5 mm); C) profemur, anterior view (2 mm); D) profemur, ventrolateral view (0.5 mm). 1996 THE GENUS CURICTA 111 parallel, longitudinal carinae present, slightly elevated. Metasternum tomentose. Parasterna of last abdominal segment ending in large processes (Fig. 26B) which extend subequally to posterior margin of operculum. Procoxae 0.54X profemoral length. Two profemoral teeth; distance from base of profemur to tip of antero ventral profemoral tooth, 0.52 X total femoral length (Fig. 26C). Profemur moderately arched beyond profemoral teeth. Profemoral sulcus with row of tubercles on distal third; these becoming more closely spaced distally except for gap occurring near junction with protibia (Fig. 26D); no sulcal teeth distally. Metafemora reaching almost halfway onto abdominal sternum 6 when extended posteriorly. Parameres; species known only from female holotype. Female genitalia: Gp2 dorsoventrally flattened (Fig. 13B). Diagnosis. This species’ very long respiratory siphons (0.86 X body length), long profemora, protibial medial annulus, prominent carinae on the vertex and pronotal ridges, and absence of profemoral sulcal teeth are, in combination, diagnostic. These diagnostic characters plus the absence of profemoral sulcal teeth in decarloi distin- guish this species from the other long-legged species found along Brazil’s coast (C. volxemi and doesburgi). C. decarloi is very close to C. carinata. Both species have dark medial protibial annuli, a prominent carina on the vertex, carinate pronotal ridges, and carinae on the mesosterna. However, carinata' s mesosternal carinae are quite prominently elevated while decarloVs are not. In addition, carinata has evenly spaced tubercles in the profemoral sulcus while in decarloi the tubercles become clustered distally with a gap just before the end of the sulcus. Finally, decarloi has proportionally longer respiratory siphons, approximately 0.9 X body length, while the siphons of carinata females attain only 0.7 X body length. It is interesting to note that the largest female carinata, which in size equal decarloi, occur in Ecuador and Colombia, i.e., the northern part of that species’ range. Further collecting across northern South America may well uncover specimens linking the northern carinata with the coastal decarloi. If that proves to be the case then the species status of decarloi will have to be reevaluated. Distribution (Fig. 35): Tapera, Pernambuco, Brazil. Material examined. HOLOTYPE, female: Curicta decarloi Keffer. Tapera, Pernam- buco, Brazil; 4 May 1938; no collector data. Deposited in the Snow Entomological Museum, University of Kansas, Lawrence, Kansas. This species is dedicated to the memory of Dr. Jose De Carlo whose career at the Museo Argentine de Ciencias Naturales “Bernadino Rivadavia” in Buenos Aires, Argentina, was devoted to the study of the aquatic Heteroptera of South America. Curicta granulosa De Carlo Figs. 27, 36 Curicta granulosa De Carlo, 1951:411-412; Nieser, 1974:131-133. Curicta venezolana De Carlo, 47-48; 1967 (1966):35. NEW SYNONYMY. Curicta intermedia [sic], Roback and Nieser, 1974:40-41. (Misidentification) Redescription Measurements. Males: Length, 18.0-22.0; profemoral length, 4. 9-6. 2; siphons, 7.0- 15.0 Females: Length, 20.0-25.0; profemoral length, 5. 6-7. 3; siphons, 12.5-20.0 178 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Fig. 27. C. granulosa: A) dorsal habitus (5 mm); B) head and anterior prothorax, lateral view (1 mm); C) meso- and metasternum (1 mm); D) last abdominal segment, lateral view (0.5 mm); E) male operculum (0.5 mm); F) profemur, anterior view (2 mm); G) parameres (0.25 mm). 1996 THE GENUS CURICTA 179 Color. Dark brown. Lighter mottling evident, particularly on the humeral lobes and hemelytra. Abdominal dorsum dark brownish red. Abdominal sterna 3-6 with four irregular, longitudinal bands of lighter color; parasterna with one such streak but fainter and less distinct. Protibia with dark basal annulation; distinct, dark medial annulation usually present. Meso- and metathoracic legs light brown to golden. Structural characteristics. Body elongate (Fig. 27 A); length 5. 3-5. 8 X maximum width. Eye width 0.5 X interocular distance (in females a slight tendency for eye width to be slightly less than 0.5 X interocular distance). Vertex usually with pronounced mediolongitudinal carina (Fig. 27B). Lobe of antennal segment 2 usually greater than 0.5 X length of 3. Pronotum elongate; lateral length approximately 1.3-1.5X posterior width. Trans- verse sulcus incomplete. Median longitudinal sulcus deep, often becoming shallower posteriorly; complete. Median longitudinal ridges prominent, carinate. Lateral sulci deep. Lateral ridges rounded. Ridges and humeral lobes almost always densely cov- ered with granulations. Posteroventral extensions of pronotum do not meet on ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width less than length. Trident pattern distinct, carinate; longitudinal section of lateral prong, medial prong, and handle black; transverse section of lateral prong lighter, often indistinct. Hemelytral greatest width variable, occurring in anterior fourth or just anterior to base of hemelytral membranes; many granulations evident, particularly along he- melytral margins and in clavus. Metathoracic wings fully developed. Last abdominal tergum tomentose with a pair of parallel, mediolongitudinal bands of longer hairs. Prosternum in lateral view not swollen in the posterior half. Mesosternum with midline groove present; usually two prominent, wide, parallel, longitudinal, tomen- tose bands present (Fig. 27C). Parasterna of last abdominal segment ending in small digitate lobes (Fig. 27D) which extend subequally to posterior margin of male and female opercula. Male operculum. Fig. 27E. Procoxae 0.45-0.55 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of antero ventral profemoral tooth 0.48-0.54 X total profe- moral length (Fig. 27F). Profemur moderately arched beyond profemoral tooth. Pro- femoral sulcus with a sparse, single row of tubercles in distal half; sulcal teeth distally, often pronounced. Metafemora usually reach onto anterior half of abdominal sternum 6 when ex- tended posteriorly (infrequently, the metafemora of females may fail to reach the anterior margin of sternum 6 when extended). Parameres Fig. 27G. Female genitalia: Gp2 dorsoventrally flattened (Fig. 13B). Discussion. This a very distinctive species. A pronounced carina on the vertex, dark, medial protibial annulus, heavily granular dorsum, broad, hairy bands on the me- sosternum and the presence of profemoral sulcal teeth are, in combination, diagnos- tic. It should be noted, however, that one or more of these diagnostic characters may often be missing from individuals within a given granulosa population. In particular, the protibial annulus may be incomplete, faint, and, in a few individuals even absent; the mesosternal bands may be incomplete, or spotty; the carina on the vertex may not be as pronounced as figured (the carinae and bands of hair are often difficult to 180 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-^) evaluate because adhering substrate may be matting down the hair that gives the carinae some, or even most, of their form; see Discussion of Characters); and/or the amount of dorsal granulation may be reduced (the same problem with the carinae applies here; in addition certain killing and/or preserving protocols reduce hairy surficial features by matting all the hair together). All specimens of granulosa throughout its range have profemoral sulcal teeth. Rarely, one finds individuals with- in populations of granulosa that have profemoral sulcal teeth and the general facies of the other members of the population but that lack a protibial band (absent more in females than in males), and apparently also lack carinae on the vertex, bands of hair on the mesosternum, and much of any granulation on the dorsum. More com- monly just one of these four characters will be absent or not found to the degree discussed. For example, a series from Itabuna, Brazil has one female lacking both a protibial annulus and mesoternal bands. However, the specimen does have carinate pronotal ridges and granulations on the dorsum and by general facies resembles the other members of the series, so I have identified it as granulosa. The synonymy of C. venezolana De Carlo with granulosa is clear. The type for venezolana possesses all the diagnostic features for granulosa. Curiously, in his original description for venezolana, De Carlo (1960:47-48) does not compare his new species with his own previously described species, granulosa. Instead, he con- trasts venezolana with Kuitert’s carinata and peruviana using paramere shape and a perceived difference in medial protibial annuli. According to De Carlo, carinata has a well defined medial annulus on the protibia while venezolana and peruviana do not. This is incorrect. His holotype for venezolana has a well defined, albeit narrow, annulus and peruviana has a distinctive, wide annulus. In addition to De Carlo’s type, three long series of granulosa have been collected at sites in Venezuela. Like the longer series from Brazil, most of the specimens in the Venezuelan series possess all or most of the diagnostic characters for granulosa discussed above. There are three species with which granulosa might be confused. The first is carinata. Both granulosa and carinata have pronounced carinae on the vertex, dark, medial protibial annuli, similar head shapes, and dense coverings of granulations on the thoracic and hemelytral dorsa. However, carinata has narrow, elevated meso- sternal carinae while granulosa has broad, tomentose bands that are not elevated. In addition, carinata never has profemoral sulcal teeth as does granulosa. The second species that is close to granulosa is borellii. Both of these species have granulations on the dorsum and light colored meso- and metathoracic legs. However, borellii has only a slight median carina on the vertex (Fig. 28B), oblique, tomentose bands on the mesosternum (Fig. 28C), rounded pronotal ridges, and almost always lacks a dark medial annulus on the protibia. In contrast, granulosa usually has a pronounced carina on the vertex (Fig. 27B), parallel bands on the mesosternum (Fig. 27C), distinctly carinate pronotal ridges, and the protibial annulus. Also, borellii males and females are generally larger and have longer profemora than granulosa specimens. Finally, the median pronotal sulcus of borellii usually becomes quite shallow posteriorly and even obsolescent in some specimens, whereas granulosa specimens all have a deeper median sulcus which may become shallower posteriorly but not to the extent evident in borellii. The third species which may cause some confusion is C. intermedia. This is an enigmatic species which is not particularly distinctive in any way and, moreover, is 1996 THE GENUS CURICTA 181 Fig. 28. C. borellii: A) dorsal habitus (5 mm); B) head and anterior prothorax, lateral view (1 mm); C) meso- and metasternum (1 mm); D) last abdominal segment, lateral view (1 mm); E) male operculum (0.5 mm); F) profemur, anterior view (2 mm); G) parameres (0.25 mm). 182 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-1) known only from the holotype. C intermedia has neither mesosternal bands of hair nor dorsal granulations like granulosa but it does have a prominent carina on the vertex, a faint, and incomplete medial protibial annulus, and profemoral sulcal teeth. The problem here is that intermedia does not look dissimilar by general facies from those few individuals of granulosa that also are lacking the same granulosa char- acters. (Several workers have previously identified specimens of what I now rec- ognize as granulosa as intermedia. For example, Nico Nieser identified a series from Puerto Lopez, Meta, Colombia, as intermedia and then used those specimens as a template in a discussion comparing intermedia’’' with doesburgi. C. doesburgi has very large, thumblike parasternal lobes while intermedia has smaller lobes; however, like doesburgi, intermedia lacks carinate pronotal ridges and heavy granulations on the dorsum so these two species are often confused; in fact, they might be the same species — see discussion for intermedia. But Nieser’s specimens have carinate pro- notal ridges, dorsal granulations, mesosternal bands, vertex carinae, and protibial annuli so they are clearly granulosa. Similarly, H. B. Hungerford identified two specimens from Bolivia as intermedia but again, the specimens possess the diag- nostic characters for granulosa.) Thus, it may be that intermedia is simply a gran- ulosa outlier; however, given that the single intermedia specimen is not part of an identifiable granulosa series, I have chosen not to consider such a possibility. In fact, I consider the synonymy of granulosa with intermedia extremely unlikely; there are too many distinctive characters separating them (see discussion of intermedia). Distribution (Fig. 36). Brazil, Paraguay, Bolivia, Peru, Colombia, Venezuela. It is of interest to note that C. granulosa is the only South American species that has been collected widely across the northern half of South America and throughout the Am- azon Basin. Material examined. HOLOTYPE, male: Curicta granulosa De Carlo: Amazonas, Mana’os, Brazil; 1899, leg. Bicego. Deposited in the Museu de Zoologia da Univ- ersidade de Sao Paulo. ALLOTYPE, female: Curicta granulosa De Carlo: same locality, temporal, and collector data as holotype. Deposited in the Museu de Zoo- logia da Universidade de Sao Paulo. PARATYPES, two females, one male: Curicta granulosa De Carlo: 2 specimens with the same locality, temporal, and collector data as holotype; 1 specimen without any data. Deposited in the Museu de Zoologia da Universidade de Sao Paulo (1 female) and the Museo Argentino de Ciencias Naturales “Bernadino Rivadavia” (1 female, 1 male). HOLOTYPE, male: Curicta venezolana De Carlo: Sosa, Guarico, Venezuela; 6-5-1951; leg. Roze. Deposited in the Carl Drake Collection, National Museum of Natural History, Washington. Two other paratypes for this species reported to be in the NMNH and the MACN were not seen. Additional Material Examined. 153 specimens collected in January (3), February (9), March (3), May (1), June (34), July (34), August (1), September (59), October (3), November (1), and December (1); deposited in AMNH, JTPC, INPA, LACM, MACN, NMNH, and SEMC. BRAZIL. Paraiba: Vic. Joao Pessoa (Sao Phe- lip). River Jurua. Bahia: 15 km E Itabuna. Para: Rio Iriri, Cam (62°40'W, 3°50'S) Altamira (ca. 100 km S). Sao Paulo: Landerwald. Goias: Santa Isabel, Ilha do Ban- anal, Rio Araguaia. Matto Grosso: Corumba; Cuiaba; Rio Araguaia, Santa Isabel. Rondonia: Porto Velho. Acre: Espiritu, Rio Yacuma, Estanque Pto. Espirito, Sar- miento. PARAGUAY. Boqueron: Puerto Casado, along Paraguay River bank in tide holes, etc. BOLIVIA. Beni: 20 km W San Borja; Ivon Beni (only locality given). 1996 THE GENUS CURICTA 183 del Sara (province unknown). PERU. Ayachuco: Prov. L. Mar Sivia, jungle, 790 m, in stagnant pools. San Martin: 8 km ENE Tarapoto, 550 m. Huanuco: Tingo, Maria. COLOMBIA. Antioquia: 2 km W Puerto Triunfo. Meta: 16 km SW Puerto Lopez, Hacienda Mosambique, temp, ponds. VENEZUELA: Portuguesa: Guanare estado. Guarico: 15 km S Calabozo; Hato Las Lajas, cr. Calabozo; 20 km N San Rafael de Orituco. Monagas: 60 km SE Maturm. Barinas: Carrotera National, 5.1 km W of Puente Ben Paez on Rio Bocomo. Curicta borellii Montandon Figs. 28, 36 Curicta borellii Montandon, 1903:100-101. Neotype here designated. Curicta riggii De Carlo, 1951:406-408; 1967 (1966):35. NEW SYNONYMY. Curicta dureti De Carlo, 1951:408-409. NEW SYNONYMY. Curicta carinata [sic], De Carlo, 1951:409-411. (Misidentification) Curicta bachmanni De Carlo, 1956:4-5. NEW SYNONYMY. Curicta sanmartini De Carlo, 1956:5. NEW SYNONYMY. Redescription Measurements. Males: Length, 21.5-26.0; profemoral length, 6. 1-7.0; siphons, 14.0- 19.5. Females: Length, 24.0-30.0; profemoral length, 6.2-1 .y, siphons, 14.0-20.0. Color. Dark brown. Lighter mottling evident on thoracic and hemelytral dorsa. Ab- dominal dorsum dark reddish-brown. Abdominal sterna 3-6 with four lighter irreg- ular, longitudinal bands; light mottling on parasterna not quite coalescing into bands. Protibia with dark basal annulation, very infrequently with medial annulation or spot. Meso- and metathoracic legs golden. Structural characteristics. Body elongate (Fig. 28 A); length 5. 5-6. OX maximum width. Eye width 0.5 X interocular distance (but sometimes slightly greater). Vertex slightly convex to almost flat (Fig. 28B), usually with a slight medical, longitudinal Carina. Lobe of antennal segment 2 usually greater than 0.7 X the length of 3. Pronotum elongate; lateral length 1.4-1.5X posterior width. Transverse sulcus in- complete. Median longitudinal sulcus shallow, usually becoming even shallower pos- teriorly; usually obsolescent posteriorly, infrequently complete. Median longitudinal ridges rounded to subcarinate in more robust specimens; when median sulcus ob- solescent posteriorly ridges fuse posteriorly to become single rounded ridge. Lateral sulci shallow. Lateral ridges rounded. Both pairs of ridges and humeral lobes beset with small black granulations. Posteroventral extensions of pronotum do not meet on ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width less than length. Trident pattern distinct, carinate; longitudinal section of lateral prongs, medial prong, and handle black, transverse section of lateral prongs often lighter, less distinct. Hemelytral width greatest just posterior to pronotum or margins subparallel from pronotum to base of membranes; hemelytral margins and clavus beset with dark granulations. Metathoracic wings fully developed. Last abdominal tergum tomentose, often with pair of parallel, mediolongitudinal bands of longer hairs occurring medially. 184 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Prosternum in lateral view not swollen in posterior half. Mesosternum with mid- line groove present; two oblique, tomentose bands, which begin posterolaterally, anterolateral to metacoxae, and slant convergently toward midline, also present (Fig. 28C) (rarely, the carinae turn and continue anteriorly as parallel carinae). Metaster- num tomentose. Parasterna of last abdominal segment end in digitate lobes, variable in shape; slender-elongate to subtriangular (Fig. 28D); usually extending posteriorly beyond the posterior margins of the male and female opercula. Male operculum. Fig. 28D. Procoxae approximately 0.45-0.53 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth typically 0.50-0.55 X total femoral length (Fig. 28F). Profemur moderately arched beyond profemoral teeth. Profemoral sulcus with single row of tubercles usually throughout sulcus length (may be missing in proximal fourth); sulcal teeth present, often pro- nounced. Metafemora just reaching anterior margin, or reaching onto anterior fourth, of abdominal sternum 6 when extended posteriorly. (Females usually exhibit the former condition, males the latter.) Parameres. Figure 28G. Female genitalia: Gp2 dorsoventrally flattened (Fig. 13B). Discussion. The granulations on the dorsum, lack of a pronounced carina on the vertex, absence of a dark, medial, protibial annulus, and tomentose, oblique meso- sternal bands are, in combination, diagnostic for this species. C. borellii might be confused with two other species, C. granulosa and C. does- burgi. Both borellii and granulosa have granulations on the dorsum and light colored legs. However, borellii has only a slight carina on the vertex (Fig. 28B), oblique mesosternal bands (Fig. 28C) (which only rarely continue anteriorly as parallel bands), no medial protibial annuli (Fig. 28F) (with only a couple of rare exceptions), and a shallow, and usually posteriorly obsolescent, median pronotal sulcus, while granulosa has a pronounced carina on the vertex (Fig. 27B), parallel mesosternal bands (Fig. 27C), medial protibial annuli (27F), and a deep and complete median pronotal sulcus. In addition, borellii males and females are larger and have longer profemora than granulosa specimens although there is some overlap in their respec- tive measurements (see “Measurements” for both species). Both borellii and doesburgi have light colored meso- and metathoracic legs, and similar profemoral lengths and pronotal traits. They also overlap in overall length measurements although borellii specimens are, on average, larger. However, borellii has parasternal lobes that are variable in shape but rarely thumblike, while doesburgi has consistently large, thumblike parasternal lobes. In addition, doesburgi usually lacks oblique mesosternal bands. Nevertheless, small borellii and large doesburgi specimens are easily confused, particularly if the borellii specimen lacks heavy gran- ulation on the dorsum and has reduced mesosternal bands. There are three such specimens from Paraguay which I have identified as borellii because they lack the parasternal trait of doesburgi, but in all other ways they closely resemble doesburgi. Further collecting, particularly across Brazil between Paraguay and Suriname, will help illuminate the relationship between these two species. (C. grandis has a disjunct distribution with populations in northern Argentina/Paraguay and Suriname. The closeness of the Argentine and Paraguayan borellii and the Suriname doesburgi might be the result of the same processes that produced the grandis disjunction.) 1996 THE GENUS CURICTA 185 It is important to note that like C. pronotata in western Mexico, borellii exhibits a continuum of robustness. Specimens from the mountainous area of northwestern Argentina generally have broader heads and pronota. In specimens from north central Argentina and Paraguay, head and pronotal widths exhibit continuous variation: in some specimens these characters are nearly identical with those seen in specimens from higher altitudes. The history of borellii has become somewhat confused. Unfortunately, syntypes for borellii have not been located. Montandon wrote that the specimens for borellii were in his collection and in the Museum of Turin. Dr. Antonio Rolando of the Dipartimento Di Biologia Animale of the Universita De Torino has informed me that borellii is not represented in the Montandon material deposited in his university (in litt.). However, I do have a specimen determined by Montandon in 1908 to be borellii, five years after the original description. At 22 mm it is smaller than the syntype series which measured 29-30 mm in length (Montandon, 1903). The syntype series represents the upper range of lengths for other females I have examined and determined to be borellii. De Carlo (1951:399-401), apparently without ever seeing any of the borellii syn- types, came to believe that specimens from Paraguay with very swollen prosterna were borellii. However, the specimen determined by Montandon to be borellii does not have a swollen prosternum, nor does Montandon’s original description mention this very noticeable character. Curiously, Kuitert (1947:43-44, unpub. diss.) appar- ently agrees with De Carlo on the determination of borellii when he writes in his redescription of borellii, under the heading “Additional Structural Characteristics,” “The posterior portion of the prosternum prominently inflated and swollen. . .” Yet one must believe that a careful observer like Montandon would certainly have no- ticed this outstanding prosternal shape and then included the trait in his description. Furthermore, in all the specimens with the swollen prosternum, believed by De Carlo to be ''borellii," the eye width strongly tends to be greater than 0.5 X the interocular distance. Yet this dimension was described as smaller for borellii . . des yeux qui sont plus petits avec I’espace interoculaire plus de deux foix plus large que le dia- metre de Toeil,” Montandon, 1903:100). Finally, unlike Montandon’s borellii spec- imen but like De Carlo’s type for grandis, De Carlo’s specimens with the swollen prosternum do not display profemoral sulcal teeth. For these several reasons, I therefore believe that De Carlo and Kuitert were mistaken in the opinion that "borellii" specimens have a swollen prosternum. In- stead, I believe these specimens with this character should be classified as C. grandis, whose description they do in fact match (as discussed under C. grandis). The synonymy of the four De Carlo species, riggii, dureti, bachmanni, and san- martini, with borellii is clear. All four De Carlo type specimens, and the specimen determined by Montandon in 1908 to be borellii, exhibit the diagnostic characters given above. De Carlo contrasts his four species with characters that are highly variable intraspecifically such as antennal and pronotal shape, body length and color. For example, he contrasts bachmanni and dureti (1956:4) on the basis of a 2 mm difference in total length and a blackish versus light brown color difference. But a 2 mm difference in body length in all but the smallest of curictan species is well within the normal range of overall lengths and it is apparent upon studying the types that the difference in color is probably due to sediment covering these specimens. 186 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) In another example, De Carlo compares sanmartini and riggii saying (1956:5) that the two species are similar in body length, siphon length, and in characteristics of the prosternum, pronotum, and vertex but the species can be differentiated by an- tennal shape and the presence of pale longitudinal bands on the abdominal venter on riggii which are absent on sanmartini. However, inspection of the types reveals that both specimens have the abdominal bands, and the variability in antennal shape is well within the norm for curictan species. In sum, I have no hesitation about synonymizing these four De Carlo species with Curicta borellii Montandon. Finally, it should be noted that De Carlo originally confused his future sanmartini with Kuitert’s carinata (De Carlo, 1951:409-410) because Kuitert failed to mention the presence of a dark, medial protibial annulus in his original description of carinata (Kuitert, 1949:62-64). Having learned from a colleague of this carinata trait, De Carlo realized that the specimens he had determined to be carinata in 1951 were instead a new species which he described as sanmartini (De Carlo, 1956:6). Distribution (Fig. 36). Argentina, Paraguay, Brazil. Material examined. NEOTYPE, male: Curicta borellii Montandon: Paraguay, Ven- zenyi, Asuncion; no temporal or collector data. As discussed above the syntypes for C. borellii Montandon have not been located. Therefore, I here designate the 1908 Montandon determined specimen of borellii as a neotype. The specimen has two determination labels, one handwritten underneath another label which is dated 1908 and has “det. Montand.” printed and the species name, author, and date in script. This specimen is deposited in the Hungarian Natural History Museum. HOLOTYPE, female: Curicta riggii De Carlo: Salta, Argentina; leg. Daguerre; no temporal data associated with the specimen although De Carlo (1951:407) lists a date of June, 1934. Microscope slide #3014 contains one antenna. Both specimen and slide de- posited in the Museo Argentino de Ciencias Naturales “Bernadino Rivadavia,” Bue- nos Aires. De Carlo never designated an allotype for this species but one specimen loaned from the MBR has an allotype label. Someone in the museum in Buenos Aires subsequently caught the mistake and affixed a handwritten “no es tipo” label. HOLOTYPE, female: Curicta dureti De Carlo: El Naranjo, Salta, Argentina; leg. Duret. De Carlo writes in his description that the specimen was collected by Duret and Mr. Martinez (1951:409). No temporal data are associated with the specimen but De Carlo gives a date of January, 1944 in the description of the species (ibid: 409). Slide preparation #3013 of an antenna. Both specimen and slide deposited in the Museo Argentino de Ciencias Naturales “Bernadino Rivadavia,” Buenos Aires. HOLOTYPE, female, Curicta bachrnanni De Carlo: Puerto Vallemi, Concepion, Par- aguay; July, 1952; leg. Bachmann. Deposited in the Museo Argentino de Ciencias Naturales “Bernadino Rivadavia,” Buenos Aires. Another specimen bears a paratype label for this specimen and two handwritten labels reading, “no es tipo” and “no publicado.” De Carlo did not publish any record of a paratype for this specimen. HOLOTYPE, female: Curicta sanmartini De Carlo: Paso de la Patria, Corrientes, Argentina; September, 1940; no collector data. Microscope slide #3015 is of antenna. Both specimen and slide deposited in the Museo Argentino de Ciencias Naturales “Bernadino Rivadavia,” Buenos Aires. ALLOTYPE, male: Curicta sanmartini De Carlo: Villarrica, Paraguay; June, 1938; leg. Schade. Microscope slide #3016 is of antenna. Both specimen and slide deposited in the Museo Argentino de Ciencias Naturales “Bernadino Rivadavia,” Buenos Aires. PARATYPES, both male: Curicta 1996 THE GENUS CURICTA 187 sanmartini De Carlo: Villa Rica, Paraguay; June, 1938; no collector data but De Carlo attributes the specimen to F. Schade (1956:5). San Cosme, Corrientes, Argen- tina; September, 1938; leg. Wurth. Microscope slide #3017 for the Paraguay speci- men; the other specimen has a labeled slide with no identifying slide number. Both specimens and slides deposited in the Museo Argentine de Ciencias Naturales “Ber- nadino Rivadavia,” Buenos Aires. Additional material examined. 65 specimens collected in January (1), February (2), March (1), April (28), May (7), September (20), and November (1); deposited in HNHM, INSB, JTPC, LACM, MACN, and NMNH. ARGENTINA. Salta: Salta. Tucuman: Horco Mollo; S. Javier; La Aquadita. Formosa: Gran Guardia. PARA- GUAY. Asuncion: Asuncion; Laguna nr Ita. Paraguari: Caacupa; Parq. Nac. Ybycui; Lagunas - Ruta, La Rosada, Ybycui Park; 10 km E Ybycui. Guaira: Villarrica. Ar- royo Yaqua Resaii Y pacai; Aregua (provinces unknown). BRAZIL. Acre: Rio Yac- umpe, Pto. Espirito, Esfanque. Curicta volxemi (Montandon) Figs. 29, 36 Nepoidea volxemi Montandon, 1895:476-477. Curicta volxemi, Champion 1901:353; Montandon, 1903:98; Kirkaldy and Torre Bueno, 1909:202; De Carlo, 1951:395. Curicta schoutedeni Montandon, 1909a: 143-144; De Carlo, 1951:417-418. NEW SYNONYMY Curicta brasiliensis De Carlo, 1951:404-405. NEW SYNONYMY. Curicta femoralis Roback and Nieser, 1974:41-42. NEW SYNONYMY. Redescription Measurements. Males: Length, 24.0-27.0; profemoral length, 7. 6-9. 3; siphons, 18.0- 21.0. Females: Length, 24.5-30.5; profemoral length, 8. 0-9. 8; siphons, 19.0-24.5. Color. Dark brown. Lighter on thoracic and hemelytral dorsa. Abdominal dorsum red to reddish brown. Abdominal venter often with two lighter, irregular longitudinal bands on sterna 3-6; parasterna with lighter mottling. Protibia dark basally; no dark medial annulation. Meso- and metathoracic legs apparently variable: dark to light colored. Structural characteristics. Body very elongate (Fig. 29 A); total length 5. 4-5. 9 X maximum width. Eye width slightly less than to subequal to 0.5 X interocular distance. Vertex nearly flat to slightly convex; slight median longitudinal carina evident in about half of specimens. Lobe of antennal segment 2 greater than half the length of 3. Pronotum elongate; lateral length L5-1.6X posterior width. Transverse sulcus in- complete. Median longitudinal sulcus variable: complete and shallow, becoming very shallow posteriorly; or shallow anteriorly, becoming obsolescent posteriorly. Median longitudinal ridges rounded; in specimens with incomplete median sulcus median ridges fuse posteriorly to become single, broad ridge. Lateral sulci shallow. Lateral ridges rounded. Ridges and humeral lobes with very few granulations evident; these occur more frequently on narrower specimens. Posteroventral extensions of prono- tum do not meet in ventral midline at junction of pro- and mesosterna (Fig. 2C). 188 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Eig. 29. C. volxemi: A) dorsal habitus (5 mm); B) last abdominal segment, lateral view (0.5 mm); C) last abdominal segment, ventral view (1 mm); D) profemur, anterior view (2 mm); E) parameres (0.25 mm). 1996 THE GENUS CURICTA 189 Scutellum width less than length. Trident distinct: medial prong, handle, and lon- gitudinal sections of lateral prongs black. Hemelytra usually widest posterior to humeral lobes; granulations along hemely- tral margins apparent but not heavy. Metathoracic wings fully developed. Last abdominal tergum tomentose laterally; longer hairs medially form two indis- tinct, parallel mediolongitudinal bands. Prosternum in lateral view not swollen in posterior half. Mesosternum has midline groove but no carinae or bands of hairs. Metathoracic sternum tomentose. Parasterna of last abdominal segment ending in large, thumblike processes (Fig. 29B), which usually extend slightly beyond posterior margin of male and female opercula (Fig. 29C). Male operculum. Fig. 29C. Procoxae typically 0.48-0.52 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth 0.55-0.64 X total femoral length (Fig. 29D). Profemur only very slightly arched beyond profemoral teeth. Profemoral sulcus with a complete row of tubercles; two sulcal teeth distally, often prominent. Metafemora reaching onto proximal half of abdominal sternum 6 when extended posteriorly. Parameres, Figure 29E. Female genitalia: Gp2 dorsolaterally flattened (Fig. 13B). Discussion. The very elongate profemora and pronota, the protibia without a dark medial annulus, and the large, thumblike parasternal lobes are, in combination, di- agnostic for this species. Two other species share the parasternal characters with volxemi: C. tibialis and doesburgi. C. tibialis is separated from volxemi by having shorter profemora that rarely ex- ceed the lateral length of the pronotum, less elongate pronotum (lateral length/pos- terior width <1.4; volxemi >1.5), and deeper sulci and more pronounced ridges on the pronotum than volxemi. In addition, volxemi lacks the unique “step-down” notch along the lateral margin of the profemoral sulcus found in tibialis (Fig. 24C). The two species form a continuum of expression of the carina on the vertex. In tibialis the Carina can be slight to pronounced, while in volxemi the range is from slight to absent. C. doesburgi has shorter profemora that only rarely equal the lateral length of the pronotum and a pronotal length to posterior width ratio (1.4-1. 5) that is usually less than volxemfs. However, it should be noted that the larger doesburgi specimens overlap in body length with the smaller volxemi and have profemora that can attain lengths up to 1 mm longer than the lateral pronotal length, and the larger doesburgi females have profemoral lengths matching the smallest female volxemi. Also, pro- notal sculpturing in doesburgi is variable, with the sculpturing evident in volxemi also found in doesburgi. In short, there could be confusion between these two species at the junction of the larger doesburgi and smaller volxemi. C. volxemi is found in southeastern Brazil, Mato Grosso, and Meta, Colombia while doesburgi is found across northern South America from Venezuela to Brazil east of the mouth of the Amazon. Further collecting, particularly in the Amazonian interior and along Brazil’s northeastern coast will help determine the degree of intermediacy between these two species. The synonymy of C. schoutedeni, C. brasiliensis, and C. femoralis with volxemi is based on the common possession of the profemoral, pronotal, and parasternal 190 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) diagnostic characters discussed above. The type for volxemi is the smallest male but otherwise matches the other specimens I have seen. The type for schoutedeni has one of the more robust pronota of the male specimens but there are other male specimens collected from Sao Paulo that are even more robust. The types for fe- moralis are among the most slender pronotally of this species, a feature shared with specimens from Mato Grosso and Rio de Janeiro, Brazil. C. brasiliensis is in the middle of the volxemi pronotal shape distribution. In short, although there is pronotal variation in this species it is no more than that judged as within the bounds of acceptable intraspecihc variation for other species in the genus. Roback and Nieser (1974) compared iheir femoralis with five other curictan spe- cies: brasiliensis, lenti, longimanus, riggii, and montei writing that all these species have a relatively long profemur but that femoralis is distinctive because its femoral teeth are “distinctly distad of the middle of the fore femur and the process of the second antennal segment [is] shorter.” Antennae have apparently little value in the taxonomy of this group (see. Discussion of Taxonomic Characters) and distinctly distal profemoral teeth (ratio of distance to teeth to femoral length >0.55) are shared by C. tibialis, volxemi, and doesburgi, but not C. riggii which I have determined to be synonymous with C. borellii. The apparent geographic disjunction of volxemi, most specimens collected in southeastern Brazil, dind femoralis, from central Colom- bia, is somewhat mitigated by the single specimen of this species collected in Utiariti, Mato Grosso and the fact that the precise locality of the volxemi type is at present unknown (see below). I would predict that other specimens of this species will be found in the geographic interval between the Mato Grosso site and Colombia, either following an Andean arc or across the Amazonian lowlands. De Carlo compared brasiliensis with another of his curictan species, C. longi- manus, writing (De Carlo, 1951:404) that the two species have the same character- istics of the head, prothorax, and profemur but that brasiliensis is shorter in body length and siphon length. However, the lengths of both of these type specimens fall within the range determined for volxemi: brasiliensis, a male measured 27 mm and longimanus, a female, measured 29.8 mm. in body length and the two specimens measured 20.8 mm and 27 mm respectively in siphon length. The siphon measure- ment for longimanus exceeds that recorded by me for specimens of volxemi but most of the volxemi specimens have broken siphons or are missing the siphons altogether. Therefore, it is likely, in my opinion, that De Carlo’s longimanus is also a synonym of volxemi. However, I have not succeeded in attempts to secure the loan of the type for C. longimanus. (This specimen is reported to be deposited in the Fundacao In- stituto Oswaldo Cruz, Rio de Janeiro, Brazil (ibid: 404) but officials of that institution have not responded to my numerous requests for loan of the type.) Therefore, al- though I believe that C. longimanus De Carlo is probably the same as C. volxemi Montandon, I will refrain from placing it in synonymy until such time as I can study the longimanus type. Distribution (Fig. 36). Brazil: Minas Gerais, Sao Paulo, Rio de Janeiro, Rio Grand do Sul, Mato Grosso. Colombia: Meta. Material examined. HOLOTYPE, male: Curicta volxemi (Montandon). Sta. Cruz, Mexique; leg. Camille Van Volxem. No temporal data given. Locality and collector data are printed on an Institut Royal des Sciences Naturelles de Belgique label. After receiving this specimens on loan from Belgium I realized that it could not be from 1996 THE GENUS CURICTA 191 Central America as the museum label indicates. The three species of Curicta occur- ring in Mexico, scorpio, hungerfordi, and pronotata, all have proximal profemoral teeth, i.e., distance from base of profemur to tip of the anteroventral profemoral tooth <0.45 X profemoral length, while volxemVs occur at 0.56 X profemoral length. Further, the elongate profemora and thumblike parasternal lobes of volxemi seemed to place it in the Brazilian fauna. I consulted with Dr. John Polhemus (Englewood, Colorado) on this matter and he was subsequently able to discover that the collector of the volxemi type, Camille Van Volxem, had made a collecting trip to Brazil shortly before his death at the age of 27 (Deyrolle, 1875). Where Van Volxem and his two companions, MM. Van Beneden and Walter de Sely-Longchamps, collected in Brazil is still unknown. G. Schmitz, curator of Hemiptera at the Musee Royal de T Afrique Centrale in Tervuren, Belgium, has checked through the archives in his museum for information about that collecting trip and only found mention of the trip in the biography of Walter Selys-Longchamps’ father, E. de Selys-Longchamps (in litt.). However, Dr. Polhemus thinks it highly likely that Van Volxem collected at the Santa Cruz locality that is near Rio de Janeiro. Therefore, I have recorded this locality on the Distribution map for the species (Fig. 36). This specimen is deposited in the Institut Royal des Sciences Naturelles de Belgique. SYNTYPE, male: Curicta schou- tedeni Montandon. Piquete, Sao Paulo, Brazil; leg. Schouteden; no temporal data. Montandon refers to other specimens in his and Schouteden ’s collections but I have only seen this one specimen. Deposited in the Institut Royal des Sciences Naturelles de Belgique. HOLOTYPE, female: Curicta braziliensis De Carlo. Therethafroli [Therespolis?], Est. do Rio., leg. A Miranda Ribeiro; March 1918. De Carlo indicates in his description that a paratype was deposited in the Paranaense Museum. Dr. William Overal, director of this museum in Curitiba, Parana, Brazil has informed me that this type is not to be found in his museum (in litt.). HOLOTYPE, male: Curicta femoralis Roback and Nieser. Colombia: Meta, 32 km ENE Puerto Lopez, Pond at Hacienda El Viento; 10 December 1969; leg. S. S. Roback. Deposited in the Academy of Natural Sciences of Philadelphia. ALLOTYPE, female: Curicta femoralis Roback and Nieser. Colombia: Meta, Laguna Humacita nr. Las Bocas Del Guyuriba; 12 December 1969; leg. S. S. Roback. Deposited in the Academy of Natural Sciences of Philadelphia. A male and female paratype of this species, indi- cated in the original description to be in the collection of Dr. Nico Nieser, were not seen. Additional material examined. 21 specimens collected in March (1), April (8), June (2), July (3), August (1), and December (1); deposited in JTPC, MACN, MZSP, NMNH, RMNH, and SEMC. BRAZIL. Parana: Curitiba. Sao Paulo: Sto. Andre; Itaquaqueoetuba; Rio das Pedras, Palmeiras. Minas Gerais: Bello Horizonte; Rio dos Velhos. Rio de Janeiro: Manguinho. Mato Grosso: Rio Papagaio, Utiariti. Rio San Francisco (eastern Brazil, state unknown); Repressa da E. E.; E. le Moult, Mendes (state unknown); Jtann[h]aeu (state unknown). Curicta doesburgi De Carlo Figs. 30, 37 Curicta doesburgi De Carlo, 1967 (1966):33; Nieser, 1975:131-132. Curicta montei De Carlo, 1960:48-49. NEW SYNONYMY. 192 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Fig. 30. C. doeshurgi: A) dorsal habitus (5 mm); B) posterolateral metasternal processes, ventral view (0.5 mm); C) last abdominal segment lateral views (0.5 mm); D) male operculum (0.5 mm); E) profemur, anterior view (2 mm); E) parameres (0.25 mm). 1996 THE GENUS CURICTA 193 Redescription Measurements. Males: Length, 18.0-23.0; profemoral length, 5. 6-6. 9; siphons, 14.0-18.0. Females: Length, 20.0-27.0; profemoral length, 6. 2-7. 8; siphons, 14.0- 24.0. Color. Dark brown. Lighter areas evident on the thoracic and hemelytral dorsa. Abdominal dorsum reddish brown. Abdominal venter mottled with lighter areas; often with two-four lighter, irregular longitudinal bands on sterna 3-6. Protibia dark basally; no dark medial annulation. Meso- and metathoracic legs golden. Structural characteristics. Body elongate (Fig. 30A); total length 4. 9-5. 8 X maxi- mum width. Eye width typically 0.5 X interocular distance. Vertex nearly flat to convex; median longitudinal carina often apparent, infrequently prominent. Lobe of antennal segment 2 greater than half length of 3. Pronotum elongate; lateral length approximately L3-1.5X posterior width. Trans- verse sulcus incomplete. Median longitudinal sulcus variable: usually shallow ante- riorly becoming obsolescent posteriorly; or shallow and complete, usually becoming shallower posteriorly. Median longitudinal ridges rounded; in specimens with incom- plete median sulcus median ridges fuse posteriorly to become single, broad ridge which may either be gradually declivent to level of transverse sulci or continue horizontally to the posterior margin of pronotum. Lateral sulci shallow. Lateral ridges rounded. Ridges and humeral lobes with few granulations evident. Posteroventral extensions of pronotum do not meet on ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutelium width less than length. Trident varies in distinctness: pattern subcarinate to carinate, usually somewhat darker on all parts than surrounding scutelium. Hemelytra usually widest posterior to humeral lobes; granulations along hemely- tral margins present but less numerous and distinct than found in C. granulosa and C. borellii. Metathoracic wings fully developed. Last abdominal tergum tomentose laterally, longer hairs form two indistinct, par- allel mediolongitudinal bands. Prosternum in lateral view not swollen in posterior half. Mesosternum has prom- inent midline groove; often tomentose posterolaterally but no extended bands of hairs or carinae present. Metathoracic sternum tomentose; posterolateral processes variable in shape but always noticeably prolonged (Fig. 30B). Parasterna of last abdominal segment ending in large, thumblike processes (Fig. 30C) which usually extend pos- teriorly slightly beyond posterior margin of opercula. Male operculum. Fig. 30D. Procoxae typically 0.50-0.55 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth 0.52-0.58 X total profemoral length (Fig. 30E). Profemur slightly arched beyond profemoral teeth. Profemoral sulcus with a complete row of tubercles; two sulcal teeth distally, often prominent. Metafemora usually reach halfway onto abdominal sternum 6 when extended pos- teriorly. Parameres, Fig. 30F. Female genitalia: Gp2 dorsolaterally flattened (Fig. 13B). Discussion. The large, thumblike parasternal lobes, the profemora up to 1 mm longer 194 JOURNAL OF THE NEW YORK ENTOMOLOGICAE SOCIETY Vol. 104(3-4) than the lateral length of the pronotum, and the protibia without a dark medial annulus are, in combination, diagnostic for this species. C. doesburgi is close to three other species, C. volxemi, C. borellii, and C. inter- media. C. doesburgi has shorter profemora (M, 5. 6-6. 9; F, 6. 2-7. 8) that only rarely equal the lateral length of the pronotum, while volxemi has more elongate profemora (M, 7. 6-9. 3; F, 7. 8-9. 8) that always exceed the lateral pronotal length by at least 1 mm. C. doesburgVs pronotal length to posterior width ratio (1.36-1.54) is usually less than volxemVs (1.50-1.60). However, the larger doesburgi specimens overlap in body length with the smaller volxemi and have profemora that can attain lengths equal to (but not exceeding) 1 mm longer than the lateral pronotal length; larger doesburgi females have profemoral lengths matching the smallest female volxemi. Also, pronotal sculpturing in doesburgi and volxemi is very similar. In short, there could be confusion between these two species at the junction of the larger doesburgi and smaller volxemi. C. volxemi is found in southeastern Brazil, Mato Grosso, and Meta, Colombia while doesburgi is found across northern South America from Ven- ezuela to Brazil east of the mouth of the Amazon. Further collecting, particularly in the Amazonian interior and along Brazil’s northeastern coast will help determine the degree of intermediacy between these two species. C. doesburgi and borellii specimens can also be very difficult to separate in the interval where their body measurements overlap (doesburgi: M, 18.0-23.0; F, 20.0- 27.0; borellii: M, 21.5-26.0; F, 24.0-30.0). Both doesburgi and borellii have light colored meso- and metathoracic legs and similar profemoral lengths and pronotal traits. However, borellii has parasternal lobes that are variable in shape but rarely thumblike (Fig. 28D), while doesburgi has consistently large, thumblike parasternal lobes (Fig. 30C). In addition, doesburgi lacks the oblique mesosternal bands of borellii, although many doesburgi specimens are tomentose in the posterolateral corners of the mesosternum, a condition which resembles that found in borellii spec- imens with reduced oblique bands. However, large doesburgi and small borellii spec- imens are easily confused particularly if the borellii specimen lacks heavy granula- tion on the dorsum and has reduced mesosternal bands. There are three such spec- imens from Paraguay that I have identified as borellii because they lack the para- sternal trait of doesburgi but in all other ways they closely resemble doesburgi. It is worth noting that C. grandis has a disjunct distribution with populations in north- ern Argentina/Paraguay and Suriname. The closeness of the Argentine and Para- guayan borellii and the Suriname doesburgi might be the result of the same processes that produced the grandis disjunction. Further collecting, particularly across Brazil, between Suriname and Paraguay, is needed to help understand the relationship be- tween these two species. C. intermedia is known from only a single specimen; the relationship between this specimen and C. doesburgi is examined in the discussion of intermedia. Mention should also be made of the possible synonymy of doesburgi with C. lend De Carlo. The type specimen for C. lend was not seen for this review, but a single specimen from Manguinhos, Brazil appears to match De Carlo’s original description (see “Comments” for lend). Interestingly, this lenti-\\\sQ specimen has the thumblike parasternal processes of volxemi and doesburgi and resembles by general facies the more pronotally robust specimens of doesburgi from Suriname. Further collecting is clearly needed to determine the relationship between these two species. 1996 THE GENUS CURICTA 195 The synonymy of C. montei with doesburgi is clear. C. montei has one of the more elongate pronota of the doesburgi specimens but the difference is within the bounds of pronotal differences found in other species, e.g., C. pronotata (Fig. 4A). Otherwise montei exhibits all the doesburgi traits. The montei holotype is one of four specimens of doesburgi from Brazil (Fig. 37). All four specimens are at the upper end of the measurement distribution for doesburgi. It would be interesting to see if further collecting between the mouth of the Amazon and the state of Pernam- buco (the current easternmost locality for doesburgi) produces specimens that are consistently on the upper end of the measurement distribution for doesburgi. Also, as noted above, the larger specimens of doesburgi including these Brazilian speci- mens, approach the lower end of the measurement distribution for volxemi. Thus, collecting for doesburgi and volxemi along Brazil’s eastern and northeastern coast will further our understanding of the separation of these two species. Finally, mention should be made of the excellent monograph by Dr. Nico Nieser on the Nepomorpha of Suriname (1975) which contains detailed information con- cerning collecting localities for C. doesburgi. Distribution (Fig. 37). Venezuela: Guarico. Guyana: East Demerara. Suriname: Su- riname. Brazil: Maranhao, Ceara, Pernambuco. Material examined. HOLOTYPE, male: Curicta doesburgi De Carlo. Suriname, leg. P. H. V. Doesburg, Jr. Temporal data given as 1014, 19-. Deposited in Rijksmuseum Van Natuurlijke Historie. ALLOTYPE, female: Curicta doesburgi De Carlo. Same data as holotype. Deposited in Rijksmuseum Van Natuurlijke Historie. There is also supposed to be a male paratype of doesburgi deposited in the Museo Argentino de Ciencias Naturales “Bernadino Rivadavia” but I have not seen this specimen. Two females with doesburgi paratype labels were loaned to me from that museum but they cannot be paratypes because they were not mentioned in the original description. Accordingly, someone in Buenos Aires has affixed handwritten labels to both of these specimens saying, “no es tipo.” HOLOTYPE, male: Curicta montei De Carlo. Ceara, Brazil. No temporal or collector data. Deposited in the C. J. Drake Collection of the National Museum of Natural History, Smithsonian Institution. Additional material examined. 167 specimens collected in January (14), March (35), April (56), May (2), July (21), August (16), September (2), October (6), and No- vember (2); deposited in JTPC, MACN, NMNH, NNC, and SEMC. (Most of these specimens were collected at the Guyana locations and are deposited in SEMC.) VENEZUELA. Guarico: Camaguan. El Limon (state unknown). GUYANA: East Demerara: Georgetown Br., Guiana Botanic Gardens; Georgetown Bn, Guiana Ponds, Trenches; Lamash Conservatory, E. Coast Demerara; Plantation Ogle, E. Coast Demerara. Mazaruni -Potaro: Kangarooma B Supuruni Creek (province un- known). SURINAME: Suriname: Paramaribo; Paramaribo Bot. Gard.; Paramaribo, Munderpark; pond at Wilhelmina’s Brug; Oost-West Verfinding. Saramacca: Ver- lengde Garnizoenspad, 4 km W Saramacca River; Garnizoenspa, jarikabakreek, km 24-25. Commewijne: edge of marsh at Belwaarde plantation; Road to Moengo, 7- 8 km E Meerzorg; Road to Moengo, 7 km E Meerzerg. Road to Damburg; Tobejuba, D.A. (provinces unknown). FRENCH GUIANA, [no specific locality given]. BRA- ZIL. Maranhao: Sao Luiz, Fonte d’aqua. Ceara: [no specific locality given]. Pernam- buco: Tapera. 196 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Curicta intermedia (Martin) Fig. 31, 37 Nepoidea intermedia Martin, 1898:66-67. Curicta intermedia, Montandon, 1903:98; Kirkaldy and Torre Bueno, 1909:202; De Carlo, 1951:417. Redescription Measurements. Male holotype: Length, 19.7; profemoral length, 6.0; siphons, absent. Color. Dark brown. Lighter mottling evident on thoracic and hemelytral dorsa. Ab- dominal dorsum dark reddish brown. Abdominal venter with paler longitudinal streaks on sterna 3-7; lighter spots on parasterna. Protibia dark basally, dark area continues dorsally and dorsolaterally half way along protibia. Mesothoracic and me- tathoracic legs light brown. Structural characteristics. Body elongate (Fig. 31 A); length 5.2X maximum width. Eye width 0.5 X interocular distance. Vertex convex, with pronounced median longitudinal carina. Lobe of antennal segment 3 greater than half length of 3. Pronotum elongate; lateral length 1.4X posterior width. Transverse sulcus incom- plete. Median longitudinal sulcus complete, becoming shallower posteriorly. Median longitudinal ridges rounded. Lateral sulci shallow. Lateral ridges not prominent. Faint granulations on humeral lobes, none apparent on ridges. Posteroventral extensions of pronotum do not meet in ventral midline at junction of pro- and mesosterna (Fig. 2C). Scutellum width less than length. Trident distinct: lateral prongs and handle car- inate, dark; medial prong less carinate but dark and visible. Hemelytra widest just posterior to humeral lobes; granulations along lateral mar- gins and in clavus present but not dense. Metathoracic wings fully developed. Last abdominal tergum tomentose laterally, longer hairs form two indistinct, par- allel mediolongitudinal bands. Prosternum in lateral view not swollen in posterior half. Mesosternum has midline groove; tomentose posterolaterally but no extended bands of hair or carinae. Meta- thoracic sternum tomentose; posterolateral processes do not appear elongate (Fig. 3 IB). Parasterna of last abdominal segment ending in digitate processes (Fig. 3 1C) which apparently do not extend beyond posterior margin of operculum. The char- acters of the last abdominal segment are difficult to evaluate because this area of the specimen has been roughly handled. The connexiva have apparently been cut or broken off distally; the parasternal lobes have been abraded and are therefore missing their covering of hair; and the operculum has been pried ventrally leaving it de- formed in the posterior fourth. The operculum was therefore not illustrated. Procoxae 0.48 X profemoral length. Two profemoral teeth; distance from base of profemur to tip of anteroventral profemoral tooth 0.5 1 X total femoral length (Fig. 3 ID). Profemur slightly arched beyond profemoral teeth. Profemoral sulcus with complete row of tubercles; two sulcal teeth distally. Metafemora missing; therefore, their length relative to abdominal sternum 6 un- known. Parameres not examined (unique specimen considered to delicate for dissection). Female genitalia unkonwn. 1996 THE GENUS CURICTA 197 C D Fig. 31. C. intermedia: A) dorsal habitus (5 mm); B) posterolateral metasternal processes, ventral view (0.5 mm); C) last abdominal segment, lateral view (0.5 mm); D) profemur, anterior view (2 mm). 198 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Discussion. This species is something of an enigma. For diagnostic characters I can only say at this time that it does not really fit satisfactorily into any of the other curictan species. By size it could be confused with two other species, C. granulosa and C. doesburgi. H. B. Hungerford and Nico Nieser are two previous workers who have identified granulosa specimens as intermedia. Louis Kuitert identified several doesburgi specimens from the Guyanas as intermedia. Like granulosa, particularly the smaller specimens from Venezuela, intermedia appears to have digitate parasternal lobes and a carina on the vertex. However, granulosa specimens usually have a heavily granulated dorsum, a distinct protibial medial annulus, and parallel bands of hair on the mesosternum. C. intermedia lacks heavy dorsal granulations (although some do occur on the pronotal humeral lobes and on the hemelytra); has the dark basal protibial annulus on the dorsal surface continuing up to the half way point on the tibia but no separate, distinct annulus; and is without mesosternal bands. However, as is pointed out in the discussion of granulosa, not all granulosa specimens have all that of species’ diagnostic charac- ters. All three of these characters may be reduced, incomplete, or even absent. The protibial and mesosternal characters seem particularly prone to reduction and ab- sence. A series of specimens collected from Puerto Triunfo, Colombia by John and Dan Polhemus illustrates this problem. The three males and four females from this series all have a carina on the vertex, but not as pronounced as is usual for granulosa-, granulations on the dorsum, but not as prolific as is usual for granulosa-, faint, incomplete, or absent medial protibial annuli; and parallel mesosternal bands. None of these specimens have all the granulosa characters but neither do they resemble closely by gestalt the intermedia type. Therefore, I have identified them as granulosa. Usually, however, enough of the characters are present to make granulosa identifi- cations readily apparent. Therefore, the differences between these two species are generally distinctive enough that I think it very unlikely that they are the same species. A more likely candidate for synonymy with intermedia is doesburgi. C. intermedia shares with doesburgi a lack of dorsal granulations but has similar general facies including all measurements and related ratios. It differs from doesburgi by apparently having digitate parasternal lobes (Fig. 3 1C), rather shortened posterolateral metas- ternal processes, and a distinctive carina on the vertex. C. doesburgi has large, thumblike parasternal lobes (Fig. 30C). Metasternal processes vary in shape in does- burgi but the processes are always more elongate (Fig. 30B) than is apparent in intermedia (Fig. 3 IB). However, the difference in the metasternal character may be more apparent than real, as it appears that the posterolateral processes of intermedia are reflected dorsoposteriorly. When the intermedia holotype is seen from a postero- ventral view, the mesosternal projections appear more elongate and not dissimilar from the doesburgi condition. Carinae on the vertex do appear on doesburgi speci- mens but only rarely do they approach the prominence found on intermedia. In sum, further collecting is needed in Colombia, and probably throughout north- ern South America, to help solve the puzzle of intermedia. We need to know the stability of the vertex and metasternal characters found on intermedia. Carinae on the vertex can vary in their prominence in species of Curicta, and the metasternal processes have been used sparingly in this review because of their plasticity. (Roback and Nieser, 1974, also found metasternal processes “too variable to be of much use” 1996 THE GENUS CURICTA 199 [p. 41] in curictan taxonomy.) Thus, the distinctions between intermedia and does- burgi, based on these characters, should not be considered strong hypotheses of species difference. There is one female specimen from Colombia, identified by Louis Kuitert as in- termedia, that bears mentioning. This specimen lacks dark protibial medial annuli, mesosternal carinae, and granulations on the dorsum so it probably is not granulosa. Further, it has small parasternal processes so it is unlikely that it belongs in does- burgi. However, it also lacks a carina on the vertex like the holotype for intermedia. Nevertheless, since it cannot be definitively placed in either granulosa or doesburgi and because it was collected in Colombia, I am letting Kuitert’s determination stand. The measurements for this specimen are as follow: length, 21.8; profemoral length, 5.9; siphon length, 16.0. Distribution (Fig. 37). Colombia. (Listed in original description as Parzudaki, Co- lombia.) Material examined. HOLOTYPE, male: Curicta intermedia (Martin). Colombia. There is no other locality, temporal, or collector data associated with the specimen. In Martin’s description he lists the specimen’s data as follows: “Columbie, Parzu- daki, 1840.” Additional material examined. 1 specimen from NMNH with the following data: Armero Tolima, Colombia; 1939; Coll 8 No 95; leg. F. L. Gallego M. (See above for discussion of this specimen.) Species Not Seen Curicta longimanus De Carlo Fig. 37 Curicta longimanus De Carlo, 1951:402—404. (Original description and notes trans- lated from the Spanish; all figures refer to the original text; my comments follow the translation.) ''Description: Female. Width of the interocular distance plus eyes 2.6; vertex without median longitudinal carina and not too convex; antennae (Fig. 22). Prothorax long, narrow, slightly robust. Pronotum (Fig. 21) anterior to transverse sulci slightly narrower than humeral lobes; length along midline 5.6; greatest width anteriorly 3.6, posteriorly 4.7; anterior margin angular, but a little rounded apically; anterior to the transverse sulci the prothorax is a little convex, with the longitudinal sulci visibly distinct. Prosternum posteriorly almost flat, very little convex. Meso- sternum almost flat, with the median groove well defined. Scutellum as in C. tibialis, but only slightly carinate. Abdomen narrow, measuring at the level of the hemelytral membrane apex [base?] 4.9. Profemora markedly long, robust (Fig. 23), 10 mm in length, with the teeth located just distal to the middle of the profemur; the upper part of both sides of the profemur without an oblique whitish band; procoxae 5.8 mm which is longer than the 4.5 mm procoxae of C. paraguayensis n. sp.; protibia a light yellowish color throughout except for a small dark area proximally; meso- and metafemora without a whitish band distally. Measurements: Holotype. Body length, 29.8; siphon length, 27. 200 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Fig. 32. Distribution map for C. bilobata, C. bonaerensis, C. pelleranoi, and C. peruviana. Distribution: Gov. Valdares, Minas, Brazil, leg. Cabral-Oliveira, August 1942. Observations: This species, so notable for the evident length of the procoxae and protibiae, can be easily distinguished from C. paraguayensis sp. nov. and C. grandis sp. nov. not only by the characteristics mentioned above but also by the flat pro- sternum and the light yellow color of the anterior legs. HOLOTYPE: In the collection of the Coleccion de Institute Osvaldo Cruz, Rfo de Janerio, Brazil [Fig. 86]. It was made available to me by Dr. Herman Lent.” Comments: I believe that this species is probably a synonym of C. volxemi. See the discussion for that species. Curicta lenti De Carlo Fig. 87 Curicta lenti De Carlo, 1 95 1 :405-406. (Original description and notes translated from the Spanish; all figures refer to the original text; my comments follow the translation.) 1996 THE GENUS CURICTA 201 Fig. 34. Distribution map for C. scorpio. 202 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Fig. 35. Distribution map for C. grandis, C. tibialis, C. carinata, and C. decarloi. Description: Male. Width of the interocular distance plus eyes 2.3; vertex without median longitudinal carina and not too convex; antennae (Fig. 51). Prothorax slightly robust. Pronotum (Fig. 24) anterior to transverse sulci slightly narrower than humeral lobes; anterior margin angular; length along midline 4.8; greatest width anteriorly 3.3, posteriorly 4.4; anterior to the transverse sulci the prothorax is only slightly convex, with the longitudinal sulci very distinct. Proster- num posteriorly almost flat, hardly elevated at all above the flanking sulci. Meso- sternum almost flat, with the median groove well defined. Scutellum moderately elevated; a median and two lateral carinae evident. Abdomen narrowing in a distinct way posteriorly, measuring at the level of the hemelytral membrane apex [base?] 4.5. Parameres Fig. 53. Profemora somewhat robust, 7.8 mm in length (Fig. 25), with the teeth located just distal to the middle of the profemur; the upper part of both sides of the profemur without an oblique whitish band; protibia and protarsus as in C. longimanus n. sp; posterior [meta-7] femur visibly thick. 1996 THE GENUS CURICTA 203 Eig. 36. Distribution map for C. granulosa, C. borellii, and C. volxemi. Measurements: Holotype (unique). Body length, 25; siphon length, 19.5. Distribution: Manguinhos, Rio de Janeiro, Brazil [Fig. 87], leg. H. S. Lopez, Feb- ruary 1942. Observations: This species is separated from C. brasiliensis sp. nov. by a pronotum which is less convex and which is almost as long in the midline as across the posterior margin; and by the shape of the antennae and parameres. HOLOTYPE: In the collection of the Coleccion de Institute Osvaldo Cruz, Rio de Janerio, Brazil. The specimen was made available to me by the distinguished hem- ipterologist Dr. Herman Lent, to whom I have the pleasure of dedicating this spe- cies.” Comments: Although De Carlo indicates that he deposited the type for this species in the Oswaldo Cruz Institute, Dr. Jose Carvalho, retired curator at the Museu Na- cional in Rio de Janeiro (now deceased), informed me that his former institution housed the lenti type (in litt.). In any event, I solicited the type from both museums. 204 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Fig. 37. Distribution map for C. doesburgi, C. intennedia, C. longimanus, and C. lend. The Museu Nacional, by mistake, sent the type for Ranatra lenti. The Oswaldo Cruz Institute did not respond to my inquiries. I have seen one specimen from Manguinhos, Brazil that is close to De Carlo’s description for lenti. The specimen, a female, has the following measurements: length, 25.1; profemoral length, 7.4; length from base of profemur to tip of antero- ventral profemoral tooth, 4.2; procoxal length, 4.0; lateral pronotal length, 6.4; pos- terior pronotal width, 4.4; and median pronotal length, 4.7. This specimen has the thumblike parasternal processes found in C. volxemi, doesburgi, and tibialis. By general facies it resembles the more pronotally robust specimens of doesburgi from Suriname. The specimen’s profemoral length to lateral pronotal length ratio is right at the cusp between volxemi and doesburgi. Therefore, I have tentatively determined this specimen to be C. lenti but with the understanding that lenti and doesburgi may prove to be synonyms. Further collecting is clearly needed from southeastern Brazil to Suriname in order to resolve this uncertainty. 1996 THE GENUS CURICTA 205 Table 1. Characters and character states used in cladistic analysis. 1. Vertex carina: (0) absent; (1) present but slight; (2) present and pronounced; (3) present as band of hair which bifurcates anteriorly 2. Pronotal median sulcus: (0) obsolescent posteriorly; (1) complete 3. Pronotal median ridges: (0) carinate; (1) rounded 4. Pronotal lateral sulci: (0) deep; (1) shallow 5. Pronotal granulations: (0) heavy; (1) light; (2) absent 6. Abdominal width: (0) greatest at base of hemelytral membranes; (1) greatest anteriorly; (2) subparallel margins 7. Last abdominal tergum: (0) carinae and bands absent; (1) parallel carinae present; (2) distinct parallel bands of hair present; (3) indistinct parallel bands of hair present 8. Prosternum: (0) not swollen; (1) swollen 9. Pronotum ventrally: (0) lobes don’t meet ventrally; (1) lobes meet midventrally 10. Mesosternal groove: (0) present; (1) absent 11. Mesosternal carinae and bands: (0) carinae and bands absent; (1) parallel carinae present; (2) parallel bands of hair present; (3) oblique bands of hair present 12. Parasternal process shape: (0) processes absent; (1) digitate; (2) broad; (3) thumblike 13. Parasternal process length: (0) processes end anterior to posterior margin of operculum; (1) processes end ca. equal to posterior margin of operculum; (2) processes end posterior to posterior margin of operculum 14. Profemoral teeth number: (0) Two; (1) One 15. Profemoral teeth position: (0) at base of femur; (1) proximal (distal to base of femur and less than halfway); (2) distal (greater than halfway) 16. Profemoral sulcus tubercles: (0) absent; (1) present throughout sulcus; (2) present in distal one-half of sulcus only 17. Profemoral sulcus teeth: (0) absent; (1) present 18. Profemoral sulcus shape: (0) linear; (1) curvilinear 19. Gp2: (0) arched dorsomedially; (1) dorsoventrally flattened 20. Ejaculatory reservoir of phallus: (0) present; (1) absent 21. Vesical rod of phallus: (0) open ventrally; (1) closed ventrally (tubular) 22. Abdominal sutures: (0) sutures delimiting sterna and parasterna don’t converge on midline of 6th segment; (1) sutures delimiting sterna and parasterna converge on midline of 6th abdominal segment CLADISTIC ANALYSIS Methods and Materials The characters and character states used in the cladistic analysis are listed in Table 1. Twenty-two characters were employed of which 13 were binary and 9 were mul- tistate. The character states for each character for all three matrices were coded numerically, e.g., 0,1, 2, 3, etc. The numbers thus assigned did not specify anything about the order or polarity of the character states (see below). Based on my study of the cladistic relationships of waterscorpion genera (Keffer, in prep.) the genera Nepa and Telmatotrephes were selected as outgroups. The full 22 characters by 18 taxa data matrix is found in Table 2. The computer program PAUP, version 3.1 (Swofford, 1993), was used to generate cladistic hypotheses. Multistate characters were specified as unordered (the Fitch 206 JOURNAL OF THE NEW YORK ENTOMOEOGICAL SOCIETY Vol. 104(3-4) Table 2. Character by taxon data matrix. 1234567891111111111222 0123456789012 Nepa 0000000000000000000000 Telmatotrephes 0100000000000000007000 C. bilobata 0111200010120122007111 C. bonaerensis 0011200100021120010111 112 2 C. pelleranoi 0111100000020120011111 C. peruviana 3111002110012012000111 C. himgerfordi 1001100111020011000111 11 2 C. pronotata 3110000111020012000111 11 2 C. Scorpio 2000001010100011001111 3 1 C. grandis 3011112110212022001111 C. tibialis J. 1010123000032021101111 21 12 C. carinata 2000002001121012001111 C. decarloi A 2000200001121022001111 C. granulosa 2000002000221212121111 1 2 C. borellii 0001010000321021101111 111 22 22 C. volxemi 0011113000032021101111 11 C. doesburgi 0011113000032021101111 11 2 C. intermedia 2011112000030021107111 parsimony option), because alternatives, e.g., Camin-Sokal parsimony, Dollo parsi- mony, and Wagner parsimony, assume knowledge, not available for this group, about how character states actually evolve. Multiple character states within any taxon were treated as polymorphisms, which they were; the alternative option, treating multiple character states as uncertainties, would have discarded those polymorphic states that increase tree lengths. Character polarities were not specified a priori. Instead, par- simony was used to assess character polarity and tree length simultaneously (ibid, p. 2). Character weightings were not employed because there was no a priori reason for believing that any of the characters chosen are more significant than any others. ACCTRAN optimization and the MULPARS option were both used. Heuristic search algorithms were utilized because of the size of the data set. All of the heuristic algorithms available in PAUP were tried. The TBR (tree-bisection-reconnection) and SPR (subtree-pruning-regrafting) heuristic algorithms both yielded the shortest length tree. 1996 THE GENUS CURICTA 207 Nepa Telmatotrephes C. bonaerensis C. pelleranoi C. bilobata C. grandis C. peruviana C. pronotata C. hungerfordi C. Scorpio C. carinata C. decarloi C. granulosa C. borellii C. intermedia C. tibialis C. volxemi C. doesburgi Fig. 38. Single most parsimonious cladogram for Curicta species. Results The analysis yielded a single most parsimonious cladogram (Fig. 38) of length 98 and Rescaled Consistency Index 0.481. The low RC indicates that there is consid- erable homoplasy in the data. Indeed, inspection of both the RC for each character (Table 3) and the list of the state changes for each tree node (Table 4) reveals that there are few unique synapomorphies, most of those being found near the base of the curictan tree. Node 2, the ingroup node, receives strong support from seven unique synapo- morphies; rounded pronotal ridges (3-1); shallow lateral pronotal sulci (4-1); digitate parasternal processes (12-1); profemoral teeth occurring distally (15-2); absence of an ejaculatory reservoir (20-1); tubular vesical rod (21-1); and convergence of su- tures on the 6th abdominal venter (22-1). However, character 3 reverses in the C. 208 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Table 3. Consistency index and rescaled consistency index for all characters used in cla- distic analysis. Character CI RC 1. Vertex carina 0.909 0.779 2. Pronotal median sulcus 0.889 0.667 3. Pronotal median ridges 0.750 0.600 4. Pronotal lateral sulci 0.250 0.143 5. Pronotal granulations 0.333 0.167 6. Hemelytral width 0.833 0.625 7. Last abdominal tergum 0.889 0.741 8. Prosternum 0.667 0.500 9. Pronotum ventrally 0.333 0.200 10. Mesosternal groove 0.500 0.167 1 1 . Mesosternal carinae & bands 0.750 0.375 12. Parasternal process shape 0.750 0.625 13. Parasternal process length 0.500 0.312 14. Profemoral teeth number 1.000 1.000 15. Profemoral teeth position 0.667 0.333 16. Profemoral sulcus tubercles 0.600 0.450 17. Profemoral sulcus teeth 0.500 0.400 18. Profemoral sulcus shape 1.000 1.000 19. Gp2 0.333 0.167 20. Ejaculatory reservoir 1.000 1.000 21. Vesical rod 1.000 1.000 22. Abdominal sutures 1.000 1.000 Scorpio clade and in C. hungerfordi and C. borellii. Similarly, character 4 also re- verses in the C. scorpio clade with parallel reversals occurring in C. pronotata and C tibialis. Thus, support for node 2 from these two characters is strong only in the context of the other five characters described. Node 3, consisting of C. bilobata and node 4, is supported by one unambiguous synapomorphy, a single profemoral tooth (14-1), and a single homoplasious syn- apomorphy, the absence of pronotal granulations (5-2), which is also found in C. decarloi. Node 4 includes C. bonaerensis and C pelleranoi and is diagnosed by a single unique synapomorphy, a curvilinear profemoral sulcus (18-1). Node 5 consists of nodes 6 and 9, i.e., all curictan species other than the C. bilobata clade. This node is supported by a single unambiguous synapomorphy, tubercles present throughout the length of the profemoral sulcus (16-1). Two other homoplasious synapomorphies help to diagnose the node: a slight carina on the vertex (1-1) and distinct parallel bands of hair on the last abdominal tergum (7-2). Both of these character states are also found in C. bonaerensis. Node 6, composed of C. hungerfordi and node 7, is supported by four highly homoplasious synapomorphies: swollen prosternum (8-1), also found in C. bona- erensis and C. Scorpio-, pronotal lobes meeting midventrally (9-1), also found in C. bilobata and C. scorpio-, mesosternal groove absent (10-1), also diagnosing node 12 1996 THE GENUS CURICTA 209 Table 4. List of character changes (^denotes unique synapomorphy). Node Character (Change) 2 3(0->l)*; 4(0->l)*; 12(0->2))*; 15(0->2)*; 20(0->l)*; 21(0->1)*; 22(0->l)* 3 5(0->2); 14(0->1)* 4 18(0->1)* 5 1(0->1); 7(0->2); 16(0->1*) 6 8(0->l); 9(0->l); 10(0->1)*; 15(2->1) 7 l(l->3); 16(l->2) 8 10(l->0); 12(2->1)*; 13(0->2) 9 2(l->0); 19(0->1) 10 l(l->2); 3(l->0); 4(0->l); 11(0->1) 11 13(0->1); 16(l->2) 12 10(0->1) 13 6(0->l); 13(0->2); 17(0->1) 14 5(0->l); 12(2->3)* 15 7(2->3)* C. bilobata 9(0->l); 11(0->1); 16(0->2) C. bonaerensis 1(0->0,1); 2(1->0,1); 7(0->0,2); (0->l); 13(0->1) C. pelleranoi 5(2->l); 19(0->1) C. grandis 2(1->0,1); 5(0->l); 6(0->l); ll(0->2); 15(l->2); 19(0->1) C. pronotata 4(0->l); 6(0->0,l); 7(2->0,l,2); ll(0->0,2) C. hungerfordi 2(1->0,1); 3(1->0,1); 5(0->l); 7(2->0,2) C. scorpio l(2->2,3); 7(2->l); 8(0->0,l); 9(0->l); 12(2->0); 15(2->1) C. carinata 10(0->1); 15(2->1,2) C. decarloi 5(0->2); 7(2->0) C. granulosa 6(0->0,l); ll(l->2); 15(2->1,2); 17(0->1) C. borellii 1(1->0,1); 2(0->0,l); 3(1->0,1); 6(1->1,2); 7(2->0,2); ll(0->3); 13(2->1,2); 16(1->1,2) C. intermedia l(l->2); 13(2->0) C. doesburgi 1(1->0,1,2); 2(0->0,l) C. volxemi 1(1->0,1); 2(0->0,l) C. tibialis 1(1->1,2); 2(0->0,l); 4(l->0); 6(l->2); 10(0->0,1); 11(0- >0,2) [see below] and C. tibialis', and profemoral teeth occurring proximally (15-1), also found in C. scorpio, C. granulosa and C. carinata. Node 7 includes C. pronotata and node 8 and is diagnosed by two homoplastic synapomorphies: a band of hair on the vertex (1-3) also found in C. scorpio', and tubercles present in the distal one-half of the profemoral sulcus (16-2), present also in C. bilobata, C. borellii, and diagnosing node 11 [see below]. Node 8 is composed of C. grandis and C. peruviana. One unique synapomorphy, broad parasternal processes (12-2) and one reversal, mesosternal groove present (10-0) support this node. Node 9, consisting of nodes 10 and 13, is supported by two homoplasious syna- pomorphies; pronotal median sulcus obsolescent posteriorly (2-0), also found in C. bonaerensis, C. hungerfordi, and C. grandis', and Gp2 dorsoventrally flattened (19- 1), also found in C. pelleranoi and C. grandis. 210 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Node 10 includes C. scorpio and node 11. Two homoplasious synapomorphies help to diagnose this node: pronounced carina on the vertex (1-2), also found in C. intermedia, C. tibialis, and C. doesburgi\ and parallel carinae on the mesosternum (11-1), also found in C. bilobata. In addition, two reversals support this node: car- inate pronotal median ridges (3-0), also present in C. hungerfordi and C. borellii. Node 11, consisting of C granulosa and node 12, is supported by two homopla- sious synapomorphies: parasternal processes ending equal to posterior margin of operculum (13-1), also found in C. bonaerensis and C. borellii; and tubercles found only in the distal one-half of the profemoral sulcus (16-2), also found in C bilobata, C. borellii, and diagnosing node 7 (see above). Node 12 is composed of C. carinata and C. decarloi and is diagnosed by a single homoplasious synapomorphy, mesosternal groove absent (10-1), which also diag- noses node 6 (see above) and C. tibialis. Node 13 includes C. borellii and node 14. Three homoplasious synapomorphies support this node: abdominal width greatest anteriorly (6-1), also present in C. pro- notata, C. grandis, and C. granulosa; parasternal processes ending posterior to pos- terior margin of operculum (13-2) which also supports node 8 (see above); pro- femoral sulcus teeth present (17-1), also found in C. granulosa. Node 14 consists of C. intermedia and node 15. One unique synapomorphy, thumb-like parasternal processes (12-3) and one homoplasious synapomorphy, light pronotal granulations (5-1), which is also found in C. hungerfordi and C. grandis, diagnose this node. Node 15, a trichotomy of C. tibialis, C. volxemi, and C. doesburgi, is supported by a single unique synapomorphy, indistinct bands of hair on the last abdominal tergum (7-3). In sum, there is strong, unequivocal support for the monophyly of Curicta as evidenced by the seven synapomorphies diagnosing node 2. However, for the 13 hypothesized groups within the genus, only six receive unambiguous support, i.e., C. bilobata clade (node 3); C. bonaerensis + C. pelleranoi (node 4); node 5 (all curictan species other than the C. bilobata clade); C. peruviana + C. grandis (node 8); C. intermedia clade (node 14); and C. tibialis + C. volxemi + C. doesburgi (node 15). The remaining groups are supported by homoplasious synapomorphies (paral- lelisms) and reversals. Thus, the hierarchic relationships of curictan species as hy- pothesized by this cladogram, particularly those grouped under node 5, must be considered highly provisional. Clearly, a better resolution of curictan species rela- tionships will require further study with additional characters. Discussion Although curictan species geographically form disjunct South and North American groups neither of these geographical assemblages appear to be monophyletic. Indeed, the hypothesized cladistic relationships of the South and North American species are rather complex. The three species in the basal C. bilobata clade are all South Amer- ican. Then, more distally, the North American species form clades with South Amer- ican species. Two of the three North American species, C. hungerfordi and C. pron- otata, form a clade with the South American C. peruviana and C. grandis. The third North American species, C. scorpio, forms a clade with the South American C. 1996 THE GENUS CURICTA 211 granulosa, C. carinata, and C. decarloi. The scorpio clade in turn is the sister group to a clade composed of the five remaining South American species. This cladistic hypothesis appears to present an unusual biogeographic pattern, i.e., the species of the basal bilobata clade are all found at the southern extreme of the genus’ range and two of the four species in the next most basal group, the hunger- fordi clade, are found at the northern geographical extreme. However, by combining what is known about the geological history of South, Central, and North America and the Caribbean with two aspects of the geographical distribution of extant Curicta species (there are no fossil curictans) one can construct a plausible historical scenario that is compatible with the cladistic hypothesis and biogeographic pattern. We can begin by outlining, in brief, the major geologic events that are believed to have occurred at the intersection of South, Central, and North America (Hallam, 1994; Brown and Gibson, 1983). In the Jurassic, North and South America were joined together and Central America, as we know it today, i.e., the land between the Isthmus of Tehunatepec and Colombia, did not exist. In the early Cretaceous, North and South America separated just to the south of the Yucatan peninsula. Subsequent- ly, volcanic islands appeared in the marine interval between southern Mexico and Colombia. These islands were pushed northeastward by the Farallon Plate, beginning in the mid-Cretaceous, to form Cuba, the Greater Antilles, and the islands off the coast of Venezuela. By the early Oligocene, another series of islands had been cre- ated between South and North America establishing a Central American Isthmus, albeit with marine gaps. The land corridor between South and North America was completed in the Pliocene with the emergence of the Isthmus of Panama and north- western Colombia. The aspects of curictan distributional data that may be relevant to this geologic history are two. First, curictan species are primarily either South American or North American with only two species with representation in Central America, i.e., C. carinata, a South American species that is also found in southeastern Panama, and C. Scorpio, a North American species that is also found in the northern part of Central America. Second, there are no curictan species found in the Caribbean is- lands (with the exception of one specimen of C. carinata found on Trinidad). The geologic history, distributional data, and cladistic hypothesis for Curicta can be synthesized in the following scenario. First, we must assume that curictan species were found, prior to the separation of South and North America, from Mexico to northern Argentina. The species in North America were phylogenetically related to those in South America through at least two lineages, of which the hungerfordi and Scorpio clades are extant representatives. With the divergence of South and North America, the two faunas were separated from one another. The absence of curictan species on the present day Caribbean islands suggests that there was no colonization from either South or North America of the islands that were pushed out into the Caribbean by the Farallon Plate. The presence of just two species in present day Central America can be explained as colonization events, from the north by C. Scorpio and from the south by C. carinata, subsequent to the completion of the Central American Isthmus in the Pliocene (although it is also possible that C. scor- pions dispersal southward could have occurred earlier as the Central American islands were linked in the post-Oligocene; C. carinata's dispersal northward however, would have had to wait for the emergence of the Isthmus of Panama and northwest Colom- 212 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) bia, the last geologic event completing the land bridge between South and North America). This ad hoc scenario, based on a cladistic hypothesis and distributional data for only one genus of 16 species, will need to be tested against cladograms and biogeo- graphic data for other groups of organisms, including those that can be dated from fossil remains. ACKNOWLEDGMENTS I am much indebted to J. E. McPherson and David King for their encouragement and support. J. T. Polhemus suggested this project to me and has assisted my effort in myriad ways; without his continuing and enthusiastic support, the results of this research would have been much diminished. Ivor Lansbury of the Hope Entomological Museum at Oxford University very generously provided me with unpublished notes on the genus Curicta. Carlos Peralta provided translations of the taxonomic papers of Jose De Carlo. Toby Schuh’s critical review greatly improved the final manuscript. I gratefully acknowledge the following persons and museums for the loan of specimens. The abbreviations preceding each entry have been used throughout this paper in lieu of full insti- tutional names. AMNH American Museum of Natural History — Randall T. Schuh ANSP Academy of Natural Sciences of Philadelphia — Donald Azuma ASU Arizona State University — Michael E. Douglas CAS California Academy of Sciences, San Francisco — Paul H. Arnaud, Jr. CNCIB Canadian National Collection of Insects, Biosystematics Research Institute, Research Branch, Agriculture Canada — L. Masner CNCIE Canadian National Collection of Insects, Entomology Research Institute, Research Branch, Agriculture Canada — K. G. A. Hamilton FSCA Florida State Collection of Arthropods — Frank W. Mead HNHM Hungarian Natural History Museum — Tamas Vasarhelyi INPA Instituto Nacional de Pesquisas da Amazonas — Jose Albertino Rafael ISNB Institut Royal des Sciences Naturelles de Belgique — L. Baert JTPC John T. Polhemus Collection, Englewood, Colorado LACM Los Angeles County Museum of Natural History — Charles Hogue LSU Louisiana State University, Department of Entomology — David A. Rider MACN Museo Argentino de Ciencias Naturales “Bernadino Rivadavia” — Axel Bachmann MLP Museo de la Plata, Universidad Nacional de La Plata — R. A. Ronderos MNHN Museum National D’Histoire Naturelle — D. Pluot-Sigwalt MNRJ Museu Nacional, Rio de Janeiro — Miguel A. Monne MZSP Museu de Zoologia da Universidade de Sao Paulo — E C. do Val NHMB Entomology Department, Natural History Museum, Basel — M. Brancucci NHRS Naturhistoriska Riksmusset, Stockholm — Per Lindskog NNC Nico Nieser Collection NMNH National Museum of Natural History, Washington, D.C. — R. C. Froeschner RMNH Rijksmuseum Van Natuurlijke Historie — Jan van Tol SIUC Southern Illinois University, Entomological Collection — J. E. McPherson SEMC Snow Entomological Museum, University of Kansas, Lawrence, Kansas — Robert W Brooks TXAM Texas A&M University, Department of Entomology — Joseph C. Schaffner UA University of Arizona, Department of Entomology — Floyd G. Werner UCD Bohart Museum of Entomology, University of California, Davis — Robert Schuster USU Utah State University, Department of Biology — Wilford J. Hanson 1996 THE GENUS CURICTA 213 ZMH Zoological Museum University of Helsinki — Antti Jansson LITERATURE CITED Berg, C. 1879a. Hemiptera Argentina. (Continuacion). An. Soc. Cient. 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Nepidae of the Western Hemisphere (Hemiptera). Unpubl. Ph.D. disser- tation, Univ. of Kansas, 253 pp. Kuitert, L. C. 1949a. Some new species of Nepidae (Hemiptera). J. Kans. Entomol. Soc. 22(2): 60-68. Kuitert, L. C. 1949b. Some new Ranatra from the Americas. J. Kans. Ent. Soc. 22(l):24-34. Lansbury, I. 1972. A review of the Oriental species of Ranatra Eabricius (Hemiptera-Heter- optera: Nepidae). Trans. R. Ent. Soc. Eond. 124(3):287-341. Lee, C. E. 1991. Morphological and phylogenetic studies on the True Water Bugs (Hemiptera: Heteroptera). Nature and Life 21(2): 1-1 83. Martin, J. 1898. Descriptions d’especes nouvelles de Nepidae [Hem.]. Bull. Soc. Ent. Fr. 23 (Fevrier):66-68. Mayr, E. 1942. Systematics and the Origin of Species. Columbia Univ. Press, New York, 334 pp. McPherson, J. E. and R. J. Packauskas. 1987. Life history and laboratory rearing of Nepa apiculata (Heteroptera: Nepidae), with descriptions of immature stages. Ann. Entomol. Soc. Am. 80(5):680-685. Menke, A. S. 1960. A taxonomic study of the genus Abedus StM (Hemiptera: Belostomatidae). University of California Publications in Entomology 16(8):393-440. Menke, A. S. 1979. Family NepidaeAVater Scorpions. Pages 70-75 in A. S. Menke, (ed.). The Semiaquatic and Aquatic Hemiptera of California (Heteroptera: Hemiptera). Bull Calif. Insect Surv. 21:xi + 166 pp. Montandon, A. L. 1895. Hemipteres nouveaux de la section de Hydrocorises Latr. Extr. Ann. Soc. Entomol. Belg. 39:471-477. Montandon, A. L. 1903. Hemipteres aquatiques. Notes synonymiques et geographiques, de- scription d’especes nouvelles. Bui. Soc. Rom. Sti. 12:97-121. Montandon, A. L. 1909a. Belostomatidae & Nepidae. Notes diverses et descriptions d’especes nouvelles. Bui. Soc. Rom. Sti. 18(2-4):137-147. Montandon, A. L. 1909b. Hydrocorises de I’Amerique du nord. Notes et descriptions d’especes nouvelles. Bui. Soc. Rom. Sti. 18(5-6): 180-1 91. Nieser, N. 1975. The Water Bugs (Heteroptera: Nepomorpha) of the Guyana region. Studies on the Eauna of Suriname and the other Guyanas, Vol. 16 no. 81, 308 pp. + 24 plates. Paterson, H. E. H. 1985. The recognition concept of species. In: E. S. Vrba, (ed.). Species and Speciation. Transvaal. Mus. Mongr. 4:21-29. Packauskas, R. J. and J. E. McPherson. 1986. Life history and laboratory rearing of Ranatra fusca (Heteroptera: Nepidae), with descriptions of immature stages. Ann. Entomol. Soc. Am. 79(4):566-571. Polhemus, D. A. 1988. Eamily Nepidae Eatreille, 1802. Pages 528-532 in: T. J. Henry and R. C. Eroeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Con- tinental United States. E. J. Brill, Leiden, xix + 958 pp. Polhemus, J. T. 1976. Notes on North American Nepidae. Pan-Pac. Entomol. 52(3):204-208. Roback, S. S. and N. Nieser. 1974. Aquatic Hemiptera (Heteroptera) from the Llanos of Co- lombia. Proc. Acad. Nat. Sci. Philad. 126(4):29-49. Seidenstiicker, G. 1963. Zur Afklarung von Nepa dollfusi (Heteroptera) mit 29 figuren. Rei- chenbachia 1(37):3 15-322. 1996 THE GENUS CURICTA 215 Stal, C. 1862. Nova methodus familias quasdam Hemipterorum disponendi. Ofv. K. Svensk. Vet.-Akad. Forh. 18(4): 195-2 12 (1861). Stal, C. 1865. Hemiptera Africana. 3. — Ofic. Norstedtiana Holmiae, 200 pp. Swofford, D. L. 1993. PAUP: Phylogenetic Analysis Using Parsimony, Version 3.1 Computer program distributed by the Illinois Natural History Survey, Champaign, Illinois. Tamanini, L. 1973. Priorita e sinonimia di Nepa cinerea Linneo e Nepa rubra Linneo. Regione tipica e valore delle razze europee di Nepa cinerea Linneo, 1758 (Hemiptera-Heteroptera, Nepidae). Studi Trentini Sci. Nat. (B) 50(2):222-259, 94 figs., 4 tabs. Wiley, G. O. 1922. Some notes on the biology of Curicta from Texas. Kansas Univ. Science Bull. 14(20):507-511. Wiley, G. O. 1924. On the biology of Curicta drakei Hungerford (Heteroptera, Nepidae). Entomol. News 192:324-331. Received 1 August 1996; accepted 17 March 1997. J. New York Entomol. Soc. 104(3-4):216-220, 1996 A NEW NEOTROPICAL FUNGUS GNAT (DIPTERA: SCIAROIDEA: KEROPLATIDAE) WITH MYRMECOPHAGOUS LARVAE Loic Matile Laboratoire d’Entomologie et EP 90 du CNRS, Museum national d’Histoire naturelle, 45 rue Buffon, F-75005 Paris Abstract. — The imago of Proceroplatus belluus n. sp. from Panama, a Keroplatidae with myrmecophagous larvae, is described. Other than its biology, the species is remarkable by its pectinate antennae, up to now unknown in Proceroplatus. Its taxonomic position is discussed; the species belongs to an inferred monophyletic group comprising two other species from Paraguay and Bolivia. The larvae of Keroplatidae have very contrasting trophic specializations, being either ferocious predators, killing their prey by way of toxic diffuse nets (Arach- nocampa, Macrocera and most Orfeliini), or fungicolous insects, spinning sheet-like webs to gather the spores of polyporous fungi (most Keroplatini) (see Matile, 1997, for a phylogenetic classification of the family). A phylogenetic study has shown that predation is most probably the ancestral diet of the Keroplatidae (Matile, 1997), and in the last few years, three species of Keroplatidae with ant-eating larvae have been discovered. Two of them are Oriental and belong to the genera Truplaya Edwards and Platyceridion Toilet (Kovac and Matile, in press. Chandler and Matile, in prep.); the larva of the first spins its web in bamboo internodes, that of the second in the internodes of an ant-plant (Krombein, pers. comm.). The third one is Neotropical (Panama) and belongs to Proceroplatus Edwards. The purpose of this paper is to provide a name and description for this Neotropical species, the biology of which is discussed elsewhere in this journal (Aiello and Jolivet, 1996). The larva of this very interesting species will be described in another paper. Proceroplatus is a mainly pantropical genus of Orfeliini comprising 33 described species, with a few northern representatives such as Proceroplatus elegans (Coquil- lett), which covers North America from Florida to Quebec (Laffoon, 1965). A fossil species, P. hennigi Schmallfuss is also known from the Oligocene-Miocene Domin- ican amber (Schmallfuss, 1979). The species described here differs from all the previously described species of the genus by its pectinate antennae. Nothing was known until now of the biology of any Proceroplatus’, the larvae of Proceroplatus belluus have been discovered in one of the two pouches of ant-plants of the genus Besleria (Gesneriaceae), where they prey on the ants living in the other pouch (Jo- livet, 1996; Aiello and Jolivet, 1996; Windsor and Jolivet, in press) — it is not yet known whether other species of Proceroplatus worldwide are associated with ants. MATERIAL AND METHODS In the course of the present study, I have studied 1 3 of the 1 6 described Neotrop- ical species of the genus [not seen: P. borgmerieri (Shaw), variventris (Edwards) 1996 NEW KEROPLATIDAE 217 and vitattus (Fisher)], and about twice this number of undescribed ones; they stand in the collections of the Museum national d’Histoire naturelle, Paris, or in Lane’s material kept in the Museo de Zoologia, Sao Paulo, and kindly loaned by Dr. N. Papavero. I also have examined about 20 described or undescribed species from various other biogeographical regions. The morphological terminology follows Matile (1990). The drawings have been done under the camera lucida, for the genitalia after treatment with potassium hy- droxyde (KOH). By convention, the macrochaetae of the genitalia are represented on the left side of the drawing only. Measurements were taken by means of ocular and objective micrometers. The holotype, in perfect condition, is glued at the tip of a triangular piece of cardboard; the pregenital segments and genitalia are stored in a microvial with glycerine. DESCRIPTION Proceroplatus belluus, new species (Fig 1, 2) Description (male only). — A Proceroplatus with pectinate antennae and wings strongly darkened at apex but without clear round spots (fig. 1). Length of wing: 4.4 mm. Head: occiput brown black. Three ocelli near middle of frons, the outer large, the median punctiform. Ocellar calli black. Frons dark brown. Antennae: scape and pedicel discoid, the scape black brown, the pedicel yellow. Flagellum: segments 1- 13 with long and simple pectinations bearing dense fine setae. First flagellomere yellow, the following brownish yellow, the pectinations brown, narrowly yellowed at basis. Face brownish yellow, palpi brown black, the last palpomere yellow. Thorax: prothorax, scutum, scutellum and mediotergite yellow. Pleurae and latero- tergite yellow, mesanepisternite brownish, with a group of small dorsal setae, me- sokatepisternite light brown. Laterotergite with long erect posterodorsal setae. Coxae and legs yellow, the tarsi darkened. Spurs black, the I and outer II-III minute, the inner II-III very long. Protarsus I longer than tibia (5.5:4). Wings yellow with brown spots. A costal median spot, reaching to C and M4; R4 narrowly seamed with brown, a wide apical spot, apex of M2 and M4 narrowly brownish, Culb widely seamed with brown at apex, and a weaker spot in the anal field. Costa reaching middle of section R5-M1. Subcosta short, ending in costa a little after level of rs. R4 strongly oblique, as long as costal section R1-R4. Radi- omedian fusion much shorter than stem of anterior fork (0.8: 3.2). Anal long and fine, reaching wing margin. Halteres orange yellow. Abdomen: Pattern obscured by the ciliation. Tergite I yellow, II yellow, apex slightly brownish; III brown, indistinctly yellow a little before apex; IV yellow dorsally, with a narrow postbasal brown band; V yellow, VI-VII brown, dark yellow at basis. Sternites with the same pattern as tergites. Hypopygium (Fig. 2) yellow on basal half, brown on distal half. Ninth tergite shorter than synsclerite, wider than long, concave at basis, slightly convex at apex. Cerci wide, subtriangular with rounded corners. Synsclerite simple, with a wide triangular ventral notch. Gonostyles deeply divided into two strongly sclerotized arms, the anterior regularly pointed, the posterior one wider at basis. 218 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Fig. 1. Proceroplatus belluus n. sp., male holotype, habitus. Holotype: male, “Panama, Code Prov. above El Cope, ex larva, 8 Nov. 1992 (Jo- livet, Windsor, Aiello)/Ant-eating larva on silk inside inflated Ivs of Besleria for- micaria. Adult fly 3 Jan 1984 Aiello Lot 92-87 #3.” Museum national d’Histoire naturelle, Paris. Etymology: From the Latin belluus, monster, as the larva of this species was first nicknamed by its discoverers. DISCUSSION In his very short generic description, Edwards (1925) mentioned the antennae of Proceroplatus as “much flattened, the flagellar segments deeper than long, and gen- erally articulated above the middle.” This is the condition of the males of all known species except the present one (the state of the character is attenuated in females). Pectination has appeared several times in different clades of the Keroplatinae and the Sciaroidea (see Matile, 1990:395, for a discussion of this character and its re- 1996 NEW KEROPLATIDAE 219 Fig. 2. Proceroplatus belluus n. sp., male holotype, synclerite and gonostyles, ventral view. partition), and no other imaginal character has been found to support a generic distinction between the species described here and the rest of Proceroplatus. A similar situation exists in the mycetophilid genus Metanepsia, from the Old World tropics, where the male flagellum is very similar to Proceroplatus, except for an Afrotropical species that has pectinate antennae (Matile, 1980). The two-armed structure of the male gonostyles of P. belluus is of the common Proceroplatus type, found worldwide in many species. This structure probably cor- responds to the groundplan of the genus, and is therefore of little use in assessing the phylogenetic relationships of the new species. However, the description of P. variventris Edwards, a Bolivian species known only from the female holotype (Edwards, 1931), agrees rather closely with that of P. belluus, notable differences being the antennal scape and the whole palpi, “ochreous” instead of black in the Bolivian species, and the pleural sclerites entirely yellow. The coloration of the abdomen is also different, being more strikingly band- ed. Most Proceroplatus have pictured wings with roundish apical and marginal clear spots, usually between C and R5 and between each of the posterior veins. Procer- oplatus belluus and P. variventris differ from these species by having the wing mainly clear and deprived of these round spots, while the apex of the wing is more markedly brown. A female from Paraguay shows a similar wing pattern, the main difference from the other two species being the presence of a complete transverse dark band through the posterior fork, instead of a single spot above Culb. In most Proceroplatus worldwide as well as in the Cenozoic species, the wings are orne- 220 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) merited with round clear spots which are sometimes more or less completely fused, and I have little doubt that this is the plesiomorphic state of the genus (an autapo- morphy with regards to the other genera of Orfeliini). On the basis of this deviation from the groundplan, it thus can be inferred that P. variventris, P. belluus and the Paraguayan undescribed species together form a monophyletic group. It is of course impossible to know whether the males of the two South American species have also pectinate antennae, and if the biology of these species is the same. ACKNOWLEDGMENTS I am very grateful to Drs Annette Aiello (Smithsonian Tropical Research Institute, Panama) and Pierre Jolivet, who gave me the opportunity to describe this interesting species and gen- erously deposited the holotype in the Museum national d’Histoire naturelle, Paris, and to Dr Nelson Papavero (Museu de Zoologia, Sao Paulo) for the loan of the Orfeliini of Lane’s col- lection. Thanks are heartily given to Gilbert Hodebert (Laboratoire d’Entomologie, Museum national d’Histoire naturelle, Paris) for the fine illustration of the habitus of the new species. LITERATURE CITED Aiello, A. and P. Jolivet. 1996. Myrmecophily in Keroplatidae (Diptera, Sciaroidea). J. New York Entomol. Soc. 104:226-230. Edwards, E W. 1925. British Eungus-Gnats (Diptera, Mycetophilidae), with a revised Generic Classification of the Eamily. Trans. Entomol. Soc. Lond. 1924 (1925):505-670. Edwards, E W. 1931. Die Ausbeute der deutschen Chaco Expedition 1925-6. Diptera. XXIII. Bibionidae and XXIV. Mycetophilidae. Konowia 10:74-78. Jolivet, P. 1996. Ants and Plants. An Example of Coevolution (Enlarged Edition). Backhuys, Leiden, 303 pp. Kovac, D. and L. Matile. in press. Tmplaya ferox, a new Malayan keroplatid from bamboo phytotelmata with larvae predaceous on ants (Diptera, Mycetophiloidea). Bull. Raffles Mus. Laffoon, J. 1965. Eamily Mycetophilidae (Eungivoridae): 196-229. In: A. Stone, et al. (eds.), A Catalog of the Diptera of America north of Mexico. Agric. Handb. 276, iv + 1-1696. Matile, L. 1980. Nouvelles donnees sur les Metanepsia afrotropicaux (Diptera, Mycetophili- dae). Revue fr. Entomol., N.S. 2(3): 1 19-122. Matile, L. 1990. Recherches sur la systematique et 1’ evolution des Keroplatidae (Diptera, Mycetophilidae). Mem. Mus Natl Hist. Nat., Ser. A, Zool. 148:1-682. Matile, L. 1997. Phylogeny and Evolution of the Larval Diet in the Sciaroidea (Diptera, Bi- bionomorpha) since the Mesozoic. In Grandcolas, P. (ed.). The origin of biodiversity in insects: phylogenetic tests of evolutionary scenarios. Mem. Mus. Nat. Hist. Nat., 173: 273-303. Schmallfuss, H. 1979. Preceroplatus hennigi n. sp., die erste Pilzmiicke aus dem Dominikan- ischen Bernstein (Stuttgarter Bersteinsammlung: Diptera, Mycetophiloidea, Keroplati- dae). Stuttgarter Beitr. Naturk., ser. B 49:1-9. Windsor, D. and P. Jolivet. in press. Aspects of the morphology and ecology of two Panamanian ant-plants, Hoffmannia vesciculifera (Rubiaceae) and Besleria fonnicaria (Gesneriaceae). J. Trop. Ecol., 12. Received 28 November 1996; accepted 27 March 1997. J. New York Entomol. Soc. 104(3-4):221-225, 1996 TWO NEW SPECIES OF MEGASTYLUS FROM THE NEW WORLD (HYMENOPTERA: ICHNEUMONIDAE; ORTHOCENTRINAE) David B. Wahl 3005 SW 56th Ave., Gainesville, Florida 32608-5047 phone/fax: (352) 377-6458; e-mail dbwahl@delphi.com. Abstract. — Two new species of the orthocentrine genus Megastylus, Megastylus fallax and Megastylus panamensis, are described. Megastylus panamensis constitutes the first record of this genus from the Neotropical Region. The systematics of New World Megastylus are briefly discussed. Orthocentrinae are shown to be koinobiont endoparasitoids. The orthocentrine genus Megastylus is distributed worldwide with about 30 de- scribed species. As with many ichneumonid genera, this represents only a fraction of the total number of species. In the New World, no species have been described south of the United States. Dr. Annette Aiello and her colleagues in Panama recently reared a Megastylus from a dipterous (Keroplatidae) ant-predator. It is a new species and is described here as Megastylus panamensis to complement the biological studies of the host. An unforeseen byproduct of this study is the discovery of a new species of Nearctic Megastylus from Arizona that was confused with petilus Dasch; it is described here as Megastylus fallax. Specimens examined in this study are in the American Entomological Institute (Gainesville, Florida: AEIC). Morphological terminology mostly follows Townes (1969) as modified by Wahl (1989), with the exception that the “apical transverse Carina” is referred to as the posterior transverse carina. MSI stands for the first metasomal segment. Tl, T2, SI, S2, etc., are used for the various metasomal tergites and sternites. When the lengths of the body and wing are given, the values in pa- rentheses are those of the holotype. NEW WORLD MEGASTYLUS The genus Megastylus is monophyletic, defined by the autapomorphies of: 1) an inflated scape that has the posterior margin membranous and infolded, and 2) long notauli that extend to the mesoscutal center (Wahl and Gauld, unpublished ms.). Ichneumonologists have previously been aware of its presence in the Neotropical Region, although no species have been described from there. The late Henry Townes sorted out 38 Neotropical species in the American Entomological Institute collection; they are distributed from Mexico to Chile. Ian Gauld (pers. comm.) has found 24 undescribed species in Costa Rica alone. Dasch (1992) recognized 14 Nearctic species, dividing them into the Megastylus Group (10 species) and the Dicolus Group (4 species). These are defined as having the “anterior transverse groove of propodeum” close to the metanotum in the Me- gastylus Group or separated by about 0.3 X the propodeal length (as measured from the metapostnotal posterior margin to the propodeal apex) in the Dicolus Group. Dasch’s “anterior transverse groove” is the metapostnotum (Wahl, 1985) and the 222 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Dicolus Group’s condition represents an apparent lengthening of the metapostnotum; this is apomorphic within the subfamily. The Megastylus Group is therefore non- monophyletic and should not be recognized. The name of the Dicolus Group is based upon Dicolus Forster, a junior synonym of Megastylus. Since I believe that the informal use of generic names should be reserved for genus-groups, the Dicolus Group is henceforth referred to as the insectator species-group, (insectator Forster is the type-species of Dicolus). The American Entomological Institute has hve un- described species from Argentina, Brazil, and Peru that belong to this species-group. Using Dasch’s (1992) keys to Nearctic Megastylus species, panamensis, and petilus run to the '^Megastylus'" group. In couplet 1 of the key to the species of this group, petilus is taken off by several characters, foremost of which is the elongate (7. 7-9. OX as long as wide) first flagellomere. This character, however, may well be homoplastic. Townes’ sorting of the American Entomological Institute’s Neo- tropical Megastylus segregated a group of 12 species that lack the occipital carina, most of which also have the first flagellomere elongate; /<3/to and panamensis belong to this group. Not only do these two species differ from petilus by the lack of an occipital carina, they differ in other characters as well (dimensions of the hind femur, number of bullae in vein 2m-cu, presence/absence of the posterior transverse carina). I suspect that the three species are not closely related, and the characters of the elongate first flagellomere and absent occipital carina are homoplastic. While it is generally established that orthocentrines are parasitoids of nematocer- ous Diptera (Mycetophilidae and Sciaridae; Wahl, 1990), the inference that they are koinobiont endoparasitoids has been based upon larval morphology and the biology of related taxa (Wahl, 1990). The rearing of M. panamensis provides direct confir- mation. Megastylus fallax, new species (Fig. 1) Diagnosis. This species can be recognized by the strongly convex clypeus which drops sharply away from the midline, weakly granulate to smooth mesopleuron, one bulla of vein 2m-cu of the fore wing, complete vein 2-cu of the hind wing, absence of the posterior transverse carina, and flattened SI. Female. Unknown. Male. Structure. 1. Flagellomere 1 10.0-1 0.6 X as long as wide; flagellomere 15 with numerous erect and semi-recumbent setae, erect setae about 0.5 X as long as flagellomere; 28-36 flagellomeres present. 2. Clypeus strongly convex, dropping sharply away from midline; clypeal apex weakly concave medially. 3. Occipital carina absent. 4. Mesopleuron weakly granulate to smooth, strongly shining. 5. Me- tapostnotum about 0. 1 X as long as propodeum. 6. Posterior transverse carina of propodeum absent; lateral outlines of metapleuron and pronotum as in fig. 1. 7. Ventral surface of fore coxa with basal transverse carina present. 8. Hind femur 6.2- 6.7 X as long as wide. 9. Vein 2m-cu of fore wing with one bulla. 10. Vein 2-cu of hind wing complete. 11. Lateral outline of MSI as in fig. 1; glymma of T1 absent; SI flat in lateral profile and with its apex opposite spiracle of Tl; Tl-2 strongly granulate and without rugulae. Color. As in M. petilus. Length. 3.3— 3.8 mm (3.8 mm); fore wing 2.9— 3.5 mm (3.5 mm). 1996 NEW MEGASTYLUS 223 Figs. 1-3. Lateral aspect of propodeum and MSI. 1, Megastylus fallax Wahl; 2, Megastylus panamensis Wahl; 3, Megastylus petilus Dasch. Specimens examined. Holotype 9, UNITED STATES, Arizona, Cochise Co., 10.viii.l974, H. & M. Townes [AEIC]. Condition of holotype: intact. Paratypes: 1 d, same collection data as holotype except collected 22.viii.1974; 1 d, UNITED STATES, Arizona, Gila Co., nr. Roosevelt Lake, 24.V.1947, H. & M. Townes [AEIC]. Comment. The holotype and paratypes were described as paratypes of petilus Dasch (Dasch, 1992). Etymology. From the Latin, fallax, deceitful or false, in reference to its previously cryptic identity. Megastylus panamensis, new species (Fig. 2) Diagnosis. This species can be recognized by the moderately and uniformly convex clypeus, weakly granulate mesopleuron, extensive brownish-red coloration of the mesosoma, two bullae of vein 2m-cu of the fore wing, complete vein 2-cu of the hind wing, presence of the posterior transverse carina, and convex SI. Female. Structure. 1. Flagellomere 1 about 8. OX as long as wide; flagellomere 15 with numerous erect and semi-recumbent setae, erect setae about 0.4 X as long as 224 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) flagellomere; 35 flagellomeres present. 2. Clypeus moderately convex, evenly round- ed; clypeal apex truncate medially. 4. Occipital carina absent. 5. Mesopleuron weakly granulate, shining. 5. Metapostnotum about 0. IX as long as propodeum. 6. Posterior transverse carina of propodeum present; lateral outlines of metapleuron and prono- tum as in fig. 2. 7. Ventral surface of fore coxa with weak basal transverse carina present. 8. Hind femur about 5.8 X as long as wide. 9. Vein 2m-cu of fore wing with two bullae. 10. Vein 2-cu of hind wing complete. 11. Lateral outline of MSI as in fig. 2; glymma of T1 absent; SI convex in lateral profile and with its apex apicad spiracle of Tl; Tl-2 strongly granulate and without rugulae. Color. Head with man- dible, clypeus, paraocular area (extending to 0.5 distance between antennal socket and apex of eye), margin of antennal socket, whitish; remainder of supraclypeal area, scape, pedicel, deep brownish-red; head otherwise fuscous. Mesosoma (excepting legs) brownish-red except for white of ventral pronotal margin, and fuscous of me- tapleuron, propodeum, and metathoracic venter. Fore and middle legs with coxa, trochanter, and trochantellus, white; femur, tibia, and tarsus light brownish-red except for slightly darker tint of middle tarsus. Hind leg with basal 0.5-0. 8 of coxa (margin of brownish-yellow area irregular), trochanter, anterior surface of trochantellus, basal O.I and median 0.4 of tibia, and tibial spurs, brownish-yellow; remainder of leg dark brown with diffuse longitudinal light streaking on femur. Metasoma fuscous except for brown median 0.3 of T3. Length. 5.0 mm; fore wing 3.7 mm. Male. Unknown. Type material. Holotype $, PANAMA, Code, above El Cope, 8.xi.l992, “Aiello lot 92-87, #4”, P. Jolivet — D. Windsor — A. Aiello [AEIC]. Condition of holotype: intact. Comments. M. panamensis was reared from Proceroplatus belluus Matile, a kero- platid predaceous upon ants (Matile, 1997; Aiello and Jolivet, 1997). The adult wasp emerged 31 Dec. 1992. The larva of panamensis was torn during preparation and the relationship of the hypostomal — pleurostomal process to the labial sclerite, as well the presence or absence of the mandible, is unclear. It is very similar to the larva of Megastylus sp. 1 figured in Wahl (1986); the labial sclerite appears identical. Etymology. The specific name is derived from Panama, where the specimen was collected. Megastylus petilus Dasch (Fig. 3) Megastylus petilus Dasch, 1992:134. Type: $ [AEIC]. Diagnosis. This species can be recognized by the moderately and uniformly convex clypeus, strongly granulate mesopleuron, one bulla of vein 2m-cu of the fore wing, basally incomplete vein 2-cu of the hind wing, absence of the posterior transverse carina, and convex SI. Female. Structure. 1. Flagellomere 1 9.3-1 0.8 X as long as wide; flagellomere 15 with numerous erect and semi-recumbent setae, erect setae about as long as flagel- lomere; 42-48 flagellomeres present. 2. Clypeus moderately convex, evenly rounded; clypeal apex truncate medially. 3. Occipital carina absent. 4. Mesopleuron strongly granulate. 5. Metapostnotum about 0. IX as long as propodeum. 6. Posterior trans- 1996 NEW MEGASTYLUS 225 verse carina of propodeum absent; lateral outlines of metapleuron and pronotum as in fig. 3. 7. Ventral surface of fore coxa with basal transverse carina present. 8. Hind femur 8.0-8. 3 X as long as wide. 9. Vein 2m-cu of fore wing with one bulla. 10. Vein 2-cu of hind wing basally incomplete. 1 1. Lateral outline of MSI as in fig. 3; glymma of T1 absent; SI convex in lateral profile and with its apex apicad spiracle of Tl; Tl-2 strongly granulate and without rugulae. Color. Overall color dark brown, the following brownish-yellow: malar space; clypeus; supraclypeal area; propleuron; pronotum except for ventral margin; ventral 0.5-0. 6 of mesopleuron; basal 0.5-0. 8 of hind coxa (margin of brownish-yellow area irregular); femora; fore and middle tibia and tarsi. The following white: mandible; ventral margin of pronotum; fore and middle coxae, trochanters, and trochantelli; dorsal surface of hind trochanter and trochantellus. The following brownish-yellow: hind femur and tibia except for dark brown stripe on basal 0.3 of posterior surface; T2 except for posterolateral corners; basal 0.3 of T3. Length. 4. 3-4.9 mm (4.9 mm); fore wing 3. 6-3. 8 mm (3.8 mm). Male. Structure. As in female. Color. As in female. Length. 4.4-5. 6 mm; fore wing 3. 5-4.3 mm. Specimens examined. Holotype $ , UNITED STATES, Arizona, Cochise Co., Portal, 22.viii.1974, H. & M. Townes [AEIC]. Condition of holotype: intact. Paratypes: two 9, 9 and one S, same collection data except collected 25.viii.1974, 5.ix.l974, and 24.viii.1987. ACKNOWLEDGMENTS I thank Annette Aiello and Andy Bennett for their critical comments. LITERATURE CITED Aiello, A. and P. Jolivet. 1997 (1996). Myrmecophily in Keroplatidae (Diptera: Mycetophilo- idea). J. New York Entomol. Soc. 104:226-230. Dasch, C. E. 1992. Ichneumon-flies of America north of Mexico: Part 12. Subfamilies Micro- leptinae, Helictinae, Cylloceriinae and Oxytorinae (Hymenoptera: Ichneumonidae). Mem. Am. Entomol. Inst. 52:1-470. Matile, L. 1997 (1996). A new Neotropical fungus gnat (Diptera: Sciaroidea: Keroplatidae) with myrmecophagous larvae. J. New York Entomol. Soc. 104:216-220. Townes, H. 1969. The genera of Ichneumonidae, part 1. Mem. Am. Entomol. Inst. 11:1-300. Wahl, D. B. 1985. A revision of the genus Agathilla (Hymenoptera: Ichneumonidae). Trans. Am. Entomol. Soc. 111:265-277. Wahl, D. B. 1986. Larval structures of oxytorines and their significance for the higher clas- sification of some Ichneumonidae (Hymenoptera). Syst. Entomol. 11:117-127. Wahl, D. B. 1989. A revision of Benjaminia (Hymenoptera: Ichneumonidae, Campopleginae). Syst. Entomol. 14:275-298. Wahl, D. B. 1990. A review of the mature larvae of Diplazontinae, with notes on larvae of Acaenitinae and Orthocentrinae and proposal of two new subfamilies (Insecta: Hyme- noptera, Ichneumonidae). J. Nat. Hist. 24:27-52. Received 28 November 1996; accepted 27 March 1997. NOTES AND COMMENTS J. New York Entomol. Soc. 104(3-4):226-230, 1996 MYRMECOPHILY IN KEROPLATIDAE (DIPTERA: SCIAROIDEA) The Keroplatidae, a family of the Sciaroidea (fungus gnats), are a cosmopolitan group, and, although they are encountered frequently, very little has been published on their biology. Matile (1990) revised the Arachnocampinae, Macrocerinae and Keroplatini, and included information, where known, on immature stages. Keroplatid larvae spin silk webs and are either predaceous or fungal spore feeders. The most complete account of the natural history of any predaceous member of this family can be obtained from the numerous papers on the New Zealand Glow worm, Arachnocampa luminosa (Skuse), a fungus gnat with luminous larvae (see Pugsley, 1983, 1984, for a review of the literature and ecology of the species, and Matile, 1990, for morphology and a summary of biology). The biology of the Palaearctic spore-feeder Keroplatus tipuloides Bose is fairly well known (Santini, 1982). As regards the Neotropical region, very few larvae of keroplatids have been discovered, but there are good accounts of the behavior of the predaceous larvae of Neoditomyia Lane & Stiirm (Stiirm, 1973; also Jackson, 1974, under the generic name Orfelia, and Decou, 1983, for a cave-dwelling species). Duret (1974) and Matile (1982) reported the discovery of presumably spore-feeding larvae belonging respectively to Platyroptilon Westwood, in Argentina, and Placoceratias Enderlein, in Guadeloupe, on rotten wood invaded by polyporaceous fungi, but no description of the morphol- ogy or ethology of these larvae has been given. We here present information on the larva and pupa of a carnivorous keroplatid, Proceroplatus belluus Matile, from El Cope (elevation 750 m). Republic of Panama. It is a predator of ants, as are recently discovered Oriental species of keroplatids belonging to the same tribe, but to different genera (Kovacs and Matile, in press; Matile and Chandler, in prep.; Krombein et. al., in press). On 9 November 1992, during a study of the ant-plant Besleria formicaria Nowicke (Gesneriaceae), the inflated vesicles near the base of the leaf blade of that plant were sliced open longitudinally to permit examination of the ants (Pheidole sp.), ant brood, and ant refuse deposits inside (Windsor and Jolivet, 1996). Seven of these vesicles were occupied by peculiar elongate larvae (one per vesicle) that were 1 .5-2 cm long and about 1 mm in diameter (Figs. 1, 2). The larvae were clear, except for a brownish head, and were rounded at the head end and narrowed towards the posterior end. The head bore brown mandibles, a lateral reddish spot in the position where stem- mata might be expected, and above and in front of each reddish spot, a large, clear circle representing the antennae. The head was followed by three telescoped thoracic segments and an elongate, seemingly segmentless body. Each larva rested on a strand of silk that ran the length of the vesicle and was held in position, at intervals, by perpendicular silk threads that connected it to the vesicle walls. Elongate, white, septate blobs of sticky mucus were present on some of the short silk threads. The 1996 NOTES AND COMMENTS 227 Eigs. 1-6. Figures 1-4. The fly, Proceroplatus belluus. 1. Fly larva (the black arrow points to the head end) in its natural position within the plant vesicle (sliced open longitudinally); 2. Head end of the fly larva, resting on its mucus-covered silk strand; 3. Fly pupa, suspended among silk lines; 4. Adult fly. Figures 5 and 6. The wasp. Megastylus panarnensis. 5. Wasp cocoon, with pupa within; 6. Adult wasp and empty cocoon. larvae, which appeared to have neither prolegs nor true legs, glided along the silk threads on a bed of clear slime. They reversed direction by turning the head and doubling back on themselves, on the same silk thread. Not knowing what the larvae were feeding upon, we placed the occupied vesicles into individual petri dishes inside of ZipLoc bags, numbered them, and attempted to kept the larvae moist but well ventilated, while waiting for them to present some useful clues. On 12 November, all seven larvae still were alive, but five of them had abandoned their vesicles and gone between the leaf and the petri dish or were found gliding about the dish. We returned each to its vesicle. On 14 November, larva no. 6 was consumed by a fungus that sent out white fluff all along its body. We placed the larva in water, brought it to a boil, then preserved 228 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-J) it in 80% ethanol. One by one, more larvae sucumbed to the same fate, until by 17 December only two were left. On 23 November, when there were still three larvae left (nos. 3-5), it occurred to us that the larvae might be feeding on dead ants or other dead animal matter, so we added several freshly crushed Pheidole ants taken from a potted B. formicaria. The next day, we found that the ants had been dismembered and the larvae had dark, irregular fragments in their guts. On 1 December, because the original vesicles were decomposing, we placed each larva on top of a fresh piece of B. formicaria leaf. The leaf pieces had a variety of creatures living in the water film among the trichomes on them: mites, Collembola, small worms (nematodes?), and tiny clear Crustacea. We added several freshly crushed Pheidole and some ant garbage from the vesicles of a fallen B. formicaria leaf. By the next day, all three larvae had set up fresh silk-mucus threads, their guts were full of dark brown particles, and the ants had been dismembered and compacted into mucus-covered masses. On 9 December, we added a live mosquito to each of two dishes. The next day, the two larvae in those dishes had full guts, and the third one did not. The mosquitoes had been caught on the silk and were partially dismembered and coated with mucus. It appears that the larvae are able to capture live prey, not just scavenge. By 15 December, the fly larvae were becoming opaque. On 17 December, larva no. 5 died. On 24 December larva no. 4 was consumed by the endoparasitic larva of a parasitoid wasp. We must assume that the fly larva already was parasitized when collected, as there was virtually no possibility of access by a wasp when in culture. Upon termination of feeding, the wasp larva made a pale beige, fine silk cocoon (Fig. 5), 7 mm long and 2 mm in diameter, and rounded at the ends. The cocoon was transparent enough that a white, annulated larva could be seen inside. Incor- porated into the head end of the cocoon were the remains of the fly larva. It was not possible to discern exactly when pupation took place, but on 29 December the wasp pupa began to take on color; the petiole and the sides of the abdominal seg- ments were black, and the thorax was brown. By the next day, the adult had emerged but remained inside the cocoon. A pool of amber liquid and a compressed white shed skin filled the posterior end of the coocon. On 31 December, an adult ichneu- monid wasp emerged (Fig. 6). It had an orangish thorax and clear wings with black markings; the rest of the body was black. Dr. David Wahl identified the ichneumonid as a previously undescribed species of Megastylus (subfamily Orthocentrinae); it is described as Megastylus panamensis Wahl (Wahl, 1997). Orthocentrines are pre- sumed to be koinobiont endoparasitoids of nematocerous Diptera (Wahl, 1990, 1997), and this rearing represents the first direct confirmation. Meanwhile, on 26 December, fly larva no. 3 shortened and the thorax became wider than the head. On 28 December, it pupated among the silk strands; no cocoon was made (Fig. 3). The pupa began to darken on 1 January 1993. The thorax was beige, the legs were dark gray because of the darkening setae on them, the eyes were black, and setae were beginning to show on the dorsum of abdominal segments 1-4. By 2 January, the fly appeared to be fully formed inside the pupal skin. The antennae were very broad and pectinate, the wings were dusky, the legs were black with setae, and the abdomen was clothed in black setae. The adult fly (Fig. 4) eclosed on 3 January. Dr. Loic Matile identified the fly as belonging to the sciarioid family 1996 NOTES AND COMMENTS 229 Keroplatidae, Keroplatinae, tribe Orfeliini and representing an undescribed species of Proceroplatus with pectinate antennae. He named it P. helluus, in reference to the pet name, “monster,” that we applied to the larva. All known larvae of Orfeliini are predaceous (Matile, pers. comm.). The fly specimen is in the collection of the Paris Museum, and the wasp is at the American Entomological Institute (Gainesville, Florida). Both are labelled as Aiello Lot 92-87. — Annette Aiello and Pierre Jolivet, Smithsonian Tropical Research In- stitute, Box 2072 Balboa, Ancon, Republic of Panama; and 67 Boulevard Soult, 75012 Paris, France. ACKNOWLEDGMENTS We thank Dr. Raymond Gagne, NMNH, for his help and advice, Loic Matile for identifying and describing the fly, David Wahl for identifying and describing the wasp, and the STRI Electronic Imaging Lab for preparing the photographic prints. We are grateful to Loic Matile and David Wahl for reviewing the paper and providing valuable information and references. LITERATURE CITED Decou, V. 1983. Sur la bionomie de certaines especes d’animaux terrestres qui peuplent les grottes de Cuba. Resultats. Exped. Biospeleol. Cubano-Roumaines a Cuba 4:9-17. Duret, J. P. 1974. Notas sobre el genero Platyroptilon Westwood, 1849 (Diptera, Mycetophil- idae). Rev. Soc. Entomol. Argentina 34(3-4):289-297. Jackson, J. E 1974. Goldschmidt’s dilemma resolved: Notes on the larval behavior of a new Neotropical web-spinning mycetophilid (Diptera). Am. Midi. Nat. 92(l):240-245. Krombein, K. V., B. Norden, M. M. Rickson and E R. Rickson. Ants, wasps, bees, and other domatia occupants of a Sri Lankan myrmecophyte (Hymenoptera, Diptera, Coleoptera, Lepidoptera, Psocoptera, Collembola, Pseudoscorpionida, Araneida, and Oligochaeta. In press, Smithson. Contrib. Zool. Kovac, D. and Matile, L. 1996. A new Malayan Keroplatidae (Diptera, Mycetophiloidea) from the internodes of bamboo, with larvae predaceous on ants. Senckenberg. Biol, (in press). Matile, L., 1982. Systematique, phylogenie et biogeographie des Dipteres Keroplatidae des Petites Antilles et de Trinidad. Bull. Mus. Natl Hist. Nat. Paris, 4eme sen, 4, sect. A, 1- 2:189-235. Matile, L. 1990. Recherches sur la systematique et 1’evolution des Keroplatidae (Diptera, Mycetophiloidea). Mem. Mus. Natl. Hist. Nat. Serie A. Zool. 148, Editions du Musem Paris, 682 pp. Matile, L. 1997 (1996). A new Neotropical fungus gnat (Keroplatidae, Diptera, Sciaroidea) with myrmecophagous larvae. J. New York Entomol. Soc. 104:216-220. Pugsley, C. W. 1983. Literature review of the New Zealand glowworm Arachnocampa lumi- nosa (Diptera: Keroplatidae) and related cave-dwelling Diptera. New Zealand Entomol. 7(4):419-424. Pugsley, C. W. 1984. Ecology of the New Zealand glowworm Arachnocampa luminosa (Dip- tera: Keroplatidae), in the Glowworm Cave, Waitomo. J. R. Soc. New Zealand 14(4): 387-407. Santini, L. 1982 (1979). Contribute alia conoscenza dei Micetophilidi italiani. II. Osservazioni condotte in Toscana sull’etologia di Keroplatus tipuloides Bose (Diptera, Mycetophilidae, Keroplatinae). Frustula Ent. (N.S.) 1982 2(15):151-174. Stiirm, H. 1973. Fanggespinste und Verhalten der Larven von Neoditomyia andina und N. colombiana Lane (Diptera, Mycetophilidae). Zool. Anz. 191(l-2):61-86. 230 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) Wahl, D. 1997 (1996). Two new species of Megastylus from the New World (Hymenoptera: Ichneumonidae: Orthocentrinae). J. New York Entomol. Soc. 104:221-225. Windsor, D. M. and R Jolivet. 1996. Aspects of the morphology and ecology of two Pana- manian ant-plants, Hojfniannia vesiculifera (Rubiaceae) and Besleria formicaria (Ges- neriaceae). J. Trop. Ecol. 12:835-842. Received 28 November 1996; accepted 27 March 1997. BOOK REVIEWS J. New York Entomol. Soc. 104(3— 4);23 1—235, 1996 Studies on Hemipteran Phylogeny. — C. W. Schaefer (ed.). 1996. Proceedings, Tho- mas Say Publications in Entomology. 244 pp. Entomological Society of America, Lanham, Maryland. The editor introduces this volume with the hope that included papers will stimulate further discussion of hemipteran phylogeny. I offer in the following paragraphs com- ments on some of the issues raised by the contributors. The 11 included papers were presented in a symposium at the 18th International Congress of Entomology held in Vancouver, British Columbia, Canada, in 1988. As is typical of symposia at such congresses, the list of contributors is international. Hemiptera, as used in the title, is in the broad sense. There is broad general agree- ment concerning the monophyly of the Hemiptera and of the Heteroptera. My com- ments will concentrate to a great degree on the recognition of groups and interre- lationships among the classic Homoptera. If this book is about Hemiptera, there is also one thing that it is not about — cladistics. Don’t get me wrong, there are branching diagrams, some called dado- grams, and lists of characters, some called apomorphies, but the sum of the discus- sion is not cladistic. Although the term cladistics is used many times, there is little coherence of method, and certainly much doubt expressed concerning the usefulness of cladistic methods. Although this may seem odd for a group of papers on phylog- eny prepared in the late 1980s by which time the current methods were well tested and entrenched, it may seem less surprising when one considers that many of the arguments and much of the data come from paleontology. Because of the varied approaches of the contributors, I found interpretation and comparison of approaches and results difficult. Within the Hemiptera, fossil representation of groups not known in the Recent fauna appears to be greatest in the classic Homoptera. K.G.A. Hamilton (Ottawa) attempts to resolve some descrepancies in the classification of the Auchenorrhyncha through the use of Cretaceous fossils from the Santana formation of the Northeast of Brazil; many of these represent the oldest known fossils which possess bodies and appendages in addition to wings. Hamilton begins by saying, “One of the most widely used techniques for elucidating [evolutionary] relationships is cladistics, the deducing of sister groups and their common ancestors by the distribution of derived characters in modern taxa. This still remains a controversial technique.” He then proceeds to give a brief summary of the superfamilies in which Mesozoic fossil “Homoptera,” represented primarily by wings, might be placed. Hamilton draws several conclusions, primarily from study of the Santana fossils. The most general is that “Fossils frequently exhibit character states that cannot be predicted by cladistic analysis of recent forms.” The states which seem most prob- lematic, in his mind, are those that are suppressed altogether or totally transformed in modern lineages. No wonder that cladistics could not predict them. Hamilton concludes that four monophyletic “suborders” can be recognized within 232 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) the Hemiptera: Psyllomorpha ( = Sternorrhyncha), Cicadomorpha (Auchenorrhyncha less Fulgoroidea and possibly including Aleyrodoidea), Fulgoroidea, and Heterop- teroidea (or Heteropterodea). Hamilton presents branching diagrams to portray the composition and relationships of these groups. Unfortunately, he fails to include the Heteropteroidea in these diagrams, and therefore leaves unclear the position of that group within the Hemiptera. In a similar vein, D. E. Shcherbakov (Moscow) offers a paleontological view of auchenorrhynchan evolution. His fossil-based presentation has higher groups evolv- ing from other higher groups, as for example, “Bugs (Heteroptera) arose from some scytinopteroids, possibly from Paraknightiidae. ...” His discussion is so replete with such characterizations that nearly every group mentioned must be considered para- phyletic. His most compelling conclusion may be that although in the Triassic now extinct groups of Auchenorrhyncha still dominated, in the Cretaceous the fauna looked like the modern one. J. Koteja (Krakow) reviews the morphology of the scale insects, offering a list of characters that he hopes will be of value in phylogenetic work. Neither he, nor any other author, argues that the Coccoidea (or Coccinea) are not a monophyletic group. Koteja devotes most of his effort to determining what the ground plan of the scale insects should be, but little space is given to placing the scale insects in a broader phylogenetic perspective, which made it difficult for me to Judge many of his ar- guments about the polarity of characters. This paper will be particularly interesting to coccidologists. H. Derreck Blocker (Manhattan, Kansas) reviews the largely pre-cladistic literature dealing with auchenorrhynchan relationships. Blocker’s review makes it clear that there are virtually as many published schemes of higher group relationships within the “Homoptera” as their are possible topologies for those groups. Y. A. Popov and D. E. Shcherbakov (Moscow) portray evolution in the Coleor- rhyncha as evidenced by the fossil record. This paper has the trappings of being cladistic, offering a list of characters and a cladogram. Yet, all other aspects of the paper suggest classic paleontological reasoning. Some quotes may serve to make the point. Eirst, “Coleorrhyncha, regarded as a suborder of Hemiptera (sensu lato), forms a phyletic lineage somewhat parallel to but nevertheless distinct from Heteroptera, both descending independently from primitive Auchenorrhyncha Cicadomorpha.” Second, “Paleontological data confirm the auchenorrhynchous affinities of the Pe- loridiidae [ = Coleorrhyncha in part] beyond doubt, so the suggested synapomorphies of the family and Heteroptera should be reconsidered.” And, “Except for their flat- tened habitus, bugs [= Heteroptera] and Coleorrhyncha disagree in fundamental apo- morphies. . . . The evidence discussed above forces us to reject Schlee’s Heteropter- oidea [Coleorrhyncha + Heteroptera] and treat both Heteroptera and Coleorrhyncha as suborders of Hemiptera along with Auchenorrhyncha and Sternorrhyncha.” There may be merit in the conclusion of Popov and Shcherbakov that the Peloridiidae are relict in the far Southern Hemisphere. Their remaining conclusions are far less per- suasive. M. H. Sweet (College Station, Texas) offers one of the most character-rich con- tributions in his paper on the pregenital abdomen. Most of his treatment is devoted to homologizing sclerites, the first such effort, according to him, since 1893. He proposes the pleural origin of the connexival sclerites, adopting the terms “hypo- 1996 BOOK REVIEWS 233 pleurite” and “epipleurite” for the dorsal and ventral (or inner and outer) lateroter- gites, respectively. The value of such an undertaking would be to understand mor- phological change in the pregenital abdomen irrespective of the notoriously incon- stant spiracle positions, if such were actually possible. In my view Sweet’s contribution has some obvious merits and some equally ob- vious drawbacks. On the positive side, he recognizes four apparently monophyletic suborders within the Hemiptera (Sternorrhyncha, Auchennorhyncha, Coleorrhyncha, Heteroptera) on the basis of long appreciated and apparently apomorphic characters, rather than agonizing over variability in group-defining characters, especially for the Auchenorrhyncha. The drawbacks may be of two types. First, the characters he discusses are nearly all in the pregenital abdomen, hardly a complete sampling of morphological diversity in any of the groups. Second, many of his arguments for the apomorphous nature of characters are functional, and almost nowhere does he attempt to bring his concept of apomorphy into agreement with optimization of characters on a cladogram. Some of Sweet’s arguments are patently unconvincing. He notes, for example, that the pregenital abdomen in the Cicadellidae is relatively uniform in structure. In contrast, he finds the pregenital abdomen in the Cercopidae to be morphologically much more diverse, and concludes that there is merit in raising the latter group to superfamily rank, especially if the Cicadellidae are raised to superfamily through elevation of some subfamilies to family rank. Sweet observes that at least some Fulgoroidea have fields of trichobothria on certain abdominal sterna and hypopleurites, as earlier pointed out by Ossiannilsson (1978). China (1962) and Carver, Gross, and Woodward (1991) observed similar setae in the Peloridiidae. Sweet conjectures that the “trichobothria” in these two groups may be homologous with the abdominal trichobothria in pentatomomorphan Heteroptera. I would observe that trichobothria exist on many parts of the heterop- teran body, including the head, antennae, scutellum, femora, and in apparently non- homologous forms on the abdomen in several families. These well-documented ob- servations would seem to weaken Sweet’s tentative theory of abdominal trichoboth- rial homology for the Fulgoroidea, Peloridiidae, and Trichophora. Sweet argues that the ground plan of the Heteroptera has ventral spiracle bearing hypopleurites. He proposes a new infraordinal name — Aradomorpha — for the Ara- doidea (Aradidae + Termitaphididae). He justifies this on the basis of abdominal structure, saying that the “turned over” connexivum of the Leptopodomorpha and Pentatomomorpha sensu Sweet [= Trichophora] in which the hypopleurites are dor- salized and the epipleurites are infolded is “strictly homologous and constitute[s] a synapomorphy relating these infraorders. . . presumably more closely than either is related to the Aradoidea. The similarity of abdominal structure noted by Sweet in the Leptopodomorpha and Trichophora is not unique. Examination of abdominal structure in the Cimico- morpha, for example, reveals great variation, including ventral “hypopleurites” and dorsal “epipleurites” with spiracles 2-8 ventral on the hypopleurites in many Re- duviidae and Nabidae, for example, and a completely dorsalized connexivum with spiracles 2-8 ventral on the mediosternite in the Miridae and Tingidae. He does not resolve the incongruence of spiracle position in the trichophorous Pentatomomorpha 234 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3^) (ventral on the mediosternite, except in some Lygaeoidea) and Leptopodoidea (dorsal on the “hypopleurite,” except in Leotichius). Furthermore, Sweet disregards several other attributes which argue for the mono- phyly of the Pentatomomorpha sensu lato. Possibly most obvious among these is the remarkably similar structure of the pretarsus — unqique among the Heteroptera — in all recognized families in the group, the claws being large and curved with a prox- imally attached pulvillus extending nearly the length of the claw (see scanning mi- crographs in Schuh and Slater, 1995:figs. 10.5 G-I); the parempodia always exist as a single, symmetrical, setiform pair; and median dorsal and ventral arolia are absent in all life stages as far as is known. Also, uniquely among Heteroptera, the eggs of all Pentatomomorpha, including Aradoidea, have distinctive micropylar processes. By way of contrast, the pretarsus is Leptopodomorpha is unlike that of Pentatomo- morpha, most notably possessing arolia in some life stage as do all “lower” Het- eroptera and lacking pul villi; the eggs of Leptopodomorpha do not possess micro- pylar processes. Finally, the modest amount of available DNA sequence data groups Aradidae (not the Leptopodomorpha) with the Trichophora (Wheeler et al., 1993). In sum. Sweet’s arguments for the Aradomorpha are constructed in a phylogenetic vacuum and on the basis of a single character system which shows little consistency with other characters which show great constancy of form and which consistently argue for the monophyly of the Pentatomomorpha sensu lato. D. B. Thomas (Weslaco, Texas) reviews chromosome numbers in the Heteroptera, attempting to determine whether polyploidy has played a role in the evolution of the group. His survey of the literature makes it clear that the story told by existing knowledge is far from clear. The reasons can be stated as follows: 1) the sample of taxa is small, with little accounting for within-group variation; 2) the phylogenetic relationships he postulates to explain chromosome data are often questionable; and 3) his assumptions about the plesiomorphic condition in many groups are at most a guess. These are the obvious limitations of his analysis. Perhaps much more important is the problem of homology. Does simple counting of similar- appearing chromosomes actually represent a valid comparison? Thomas presents the results of some studies of chromosome volume that indicate otherwise. Furthermore, no banding patterns are known, thus, we might conclude that all the- ories of chromosome homology are outright suspect. Possibly symptomatic of the larger question is the so-called “m” chromosome, or micro-chromosome. This karyotypic feature has been used in establishing schemes of relationships within the Pentatomomorpha, yet it also occurs in such distantly related groups as the Nepo- morpha. I can only conclude from Thomas’ survey that karyology has a long way to go before it is capable of making a useful contribution to understanding mecha- nisms of diversification or establishing schemes of phylogenetic relationships within the Heteroptera. Four additional papers offer further observations, primarily on the Heteroptera, but provide little in the way of phylogenetic context. These are: P. Stys (Prague) on some groundplan characters in the Heteroptera; R. J. Wootton (Exeter, England) on the functional aspects of hemipteran wings; J. R. Aldrich (Beltsville, Maryland) on the status of knowledge of pheromones; and H. Mori (Tokyo) on coalescence of ventral nerve ganglia from an embryological perspective. 1996 BOOK REVIEWS 235 Summarizing my impressions of the papers of this volume, I would say the fol- lowing. The application of the paleontological methods flaws the conclusions of all papers dealing with fossils. There seems great reluctance on the part of the paleontologists to view classifications as being based on characters, rather than some abstract con- cept of taxa which allows one group to be descended from another. Because the Paleozoic and early Mesozoic Hemiptera are largely represented by wings, there may be little hope that the taxa can be precisely characterized. This hardly seems a justification for continuing to treat these extinct groups as ancestral (and paraphy- letic) simply because they appear early and disappear early. Two works widely cited are those of Carver et al. (1991) and Sorensen et al. (1995). The former is cited as a justification for rejecting the Homoptera as a natural group. The latter is cited as a source of evidence for that conclusion. All contributors in this volume echo the view that the classic Homoptera must be abandoned as a group, yet draw little in the way of coherent conclusions as to what the new scheme should be. Nonetheless, the desire to create a new subordinal nomenclature, even with little evidence to support it, seems to be great. This drive may be most conspicuous in the work of Sorenson et al. (1995) who coined several new names in a study using 18s rDNA data. In the present volume K.G.A. Hamilton is of a like mind, noting that the suborders should be designated with equivalent names. One might conclude from reading these papers that scientific problems can be solved simply by proposing new higher-group names. I fail to see what those problems are. Clearly, the most pressing grand scale problem in hemipteran phylogenetics is to refine the diagnoses of higher groups within the Homoptera and to form a stable scheme of interrelationships within those groups. The consistent application of the core principles of cladistics — the use of character congruence to test theories of homology and the use of outgroups to determine character polarity — would go a long way toward achieving that objective. — Randall T. Schuh, Department of En- tomology, American Museum of Natural History, New York, New York 10024. LITERATURE CITED Carver, M., G. E Gross and T. E. Woodward. 1991. Hemiptera. In: The Insects of Australia. CSIRO, Mebourne. China, W. E. 1962. South America Peloridiidae (Hemiptera-Homoptera: Coleorrhyncha). Trans. R. Ent. Soc. London 114:131-161. Ossiannilsson, E 1978. The Auchenorrhyncha (Homoptera) of Fennoscandia and Denmark. Fauna Ent. Scand., vol. 7, part 1. Scandinavian Science Press, Copehangen. Schuh, R. T. and J. A. Slater. 1995. True Bugs of the World (Hemiptera: Heteroptera). Clas- sification and Natural History. Cornell University Press, Ithaca, 336 pp. Sorensen, J. T, B. C. Campbell, R. J. Gill and J. D. Steffen-Campbell. 1995. Non-monophyly of Auchenorrhyncha (“Homoptera”), based on 18s rDNA phylogeny: eco-evolutionary and cladistic implications within pre-Heteropterodea Hemiptera (s.l.) and a proposal for new, monophyletic suborders. Pan-Pac. Ent. 71:31-59. Wheeler, W. C., R. T. Schuh and R. Bang. 1993. Cladistic relationships among higher groups of Heteroptera: congruence between morphological and molecular data sets. Ent. Scand. 24:121-137. J. New York Entomol. Soc. 104(3-4):236-239, 1996 The Butterflies of Venezuela, Part 1: Nymphalidae I (Limenitidinae, Apaturinae, Charaxinae). — Andrew F. E. Neild, Meridian Publications, Greenwich, London. 144 pp. and 32 color plates. ISBN 0 9527657 0 5. General review of the book This attractive book is the first of an ambitious series of four parts to be published (tentatively) over the next five years, covering the Nymphalidae, Papilionidae and Pieridae of Venezuela. Neild has done an admirable job combining descriptions of adult butterflies and what is known of their behavior and life histories with the excellent color plates of material from the Natural History Museum (London), pho- tographed by Bernard D’Abrera. In general, this collaboration provides a significant advance beyond D’Abrera’s indispensable picture books of the neotropical butterflies (which contain little information beyond identified photographs of specimens) for people seeking to identify Venezuelan material, and as such will be a useful addition to libraries and the shelves of serious butterfly collectors. The book is organized clearly, with three introductory sections before the main text. “A guide to the contents and use of this book” covers the meanings of terms in the species and generic accounts and in the plate legends, and offers a brief introduction on systematics, classification and nomenclature. For Neild, species cor- respond to actually or potentially reproductively isolated units (the biological species concept of Mayr, 1940), while subspecies are diagnosably different populations which do not intergrade (the phylogenetic species concept of Cracraft, 1983, and Nixon and Wheeler, 1990). In practice, very few Venezuelan species have been tested for biological isolation, and almost all are recognized on the basis of consistent differences in morphology where they occur in sympatry. Thus, Nield’s distinction between species and subspecies is rather arbitrary from a practical perspective, even in his own descriptions of new taxa (see below). “The study and collection of butterflies” helpfully includes a description of the rather daunting procedures that must be followed to obtain permits to collect and export butterflies from Venezuela. Neild reports that, “permits are not given for private collecting for personal benefit,” which implies that the main utility of this book to private individuals will be to curate collections that have already been amassed. Instructions for study of life histories and the photography of living but- terflies are also given. The section on preparing a butterfly collection contains some rather poor advice related to the aesthetic qualities of specimen preservation: Neild recommends removing abdomens of large, fatty species such as Morpho to degrease them (and reattaching them afterwards). Such procedures greatly reduce the scientific value of a specimen, since it can never be known if the abdomen originally belonged to that specimen or was scavenged from some other specimen and pasted on to replace a missing one, for aesthetic purposes (I have found Heliconius erato abdo- mens attached to H. melpomene specimens, and Lamas [1996] commented on a published generic description based on such a “glue job”). Given that non-scientific collection of butterflies is apparently not legal in Venezuela, instructions on the prettification of specimens seem superfluous and, if they lead to compromises of scientific data, inappropriate. Neild also describes homemade spreading boards that 1996 BOOK REVIEWS 237 require manipulation of insects after they are removed from the boards, to adjust their height on the pin. This procedure seems undesireable and unnecessary, given the widespread availability of commercially-built spreading boards that allow pins to be placed at the proper depth from the outset. Boards could also be built that allow this problem to be avoided. The short section on Venezuelan biogeography is supplemented by rather grainy black-and-white habitat photos, mostly of cloud forest, and by more useful political and physical maps on the front and rear endplates. Neild employs the up-to-date classification scheme of Harvey (1991) and cites other recent works that address higher-level nymphalid relationships (e.g., Otero, 1990; de Jong et al., 1996), resulting in a review of nymphalid classification above the generic level that reflects the state of current knowledge (in many cases rather scant, unfortunately). Descriptions of genera are also carefully researched, and sig- nificant revisionary works are generally cited. Some species are split from familiar inclusive genera into separate, smaller genera, such as Mesotaenia from Perisama and Fountainea from Memphis (and Memphis, in turn, split from Aenea). The issue of monophyly of these groups is not addressed. In addition to providing his own field observations, Neild often quotes at length from DeVries (1987) and other au- thors who have written on the natural history and early stages of particular groups. Sometimes, this practice seems excessive, such as the six verbatim paragraphs on Eunica from Jenkins (1990) on pp. 63-64. Species accounts contain the following subsections: range, subspecies, identifica- tion, habits and foodplants. Range succinctly indicates the distribution of the entire species, while the distributions of individual subspecies are addressed in more detail in the subspecies section. Descriptions of new subspecies (discussed further below) are also presented here. The identification section is usually the most extensive, describing wing-pattern and other features that allow discrimination of subspecies and of similar species from one another. The diagnostic characters of the various taxa are described in discursive paragraphs which may be rather long — the notes on identification of Memphis arachne run for almost an entire page. Keys are provided for some but not all confusing groups. The habits include altitudinal range, preferred haunts, and behavior. Many larval foodplant records are drawn from DeVries (1987), while others were contributed by Venezuelan colleagues. The plates are excellent, reproduced at 90% of life size. Many holotype, paratype and syntype specimens from the Natural History Museum in London are figured, some for the first time. Illustration of types is a nice feature because it provides an authoritative view of the specimen with which a particular name is associated. In some instances, the verso and recto figures of particular species represent different specimens, which is slightly confusing (especially when they are paratypes of new taxa, such as Memphis maria Pyrcz and Neild). The figures are numbered consec- utively, and refer to numbers given at the top of text pages, making reference be- tween the two simple. Locality data for figured specimens are presented in an ap- pendix. Also included are a checklist, a glossary of terms, and a quite extensive bibliography. The index is a bit unusual, in that it refers primarily not to page numbers but to figure numbers. However, once this is realized, it is easy to use. There is also a gazetteer of some common collecting localities, related by grid quad- rants to the map in the front endplate. 238 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 104(3-4) Descriptions of new taxa In this book, Neild and coauthors describe 24 new subspecies and two new spe- cies. In my view, this is extremely inappropriate, for a number of reasons. First, it is unclear whether or not these descriptions were subject to peer review. Second, one species {Memphis viloriae Pyrcz and Neild) and three subspecies are described from single specimens, resulting in the third problem, which is that many of the descriptions are not clearly based on consistent differences between demonstrably distinct taxa. The genitalia of only one of the new taxa are illustrated. Fourth, three of the holotypes reside in private collections, with no indication of intentions to deposit them in a permanent and publicly accessible museum. Fifth (and perhaps most important), it is not clear to what extent Neild and colleagues have sought collections outside Britain and Venezuela, and in particular in the United States, to examine additional material. While this review is not the place for a thorough investigation of these problems, I make the following observations as an indication of the importance of thorough exploration of known collections before describing new taxa: I have seen seven specimens of the “new” subspecies Adelpha olynthia pyrczi in the Smithsonian, and five additional specimens in the American Museum of Natural History (three of which are identified as Adelpha olynthia inachia Fruhstorfer, a name not mentioned in Neild’s book). It is ironic that although Neild dedicates the book to (among others) William Beebe, he has not examined Beebe’s material from Rancho Grande, which is in the AMNH (nine of Neild’s new subspecies occur in the Rancho Grande region). Furthermore, I have found at least one species of Adelpha in the Smithsonian, evi- dently collected in Venezuela, that Neild does not mention or illustrate in the book. That I was able to find these records with relatively little effort or expertise in the groups concerned is a clear indication of Neild’s need to study material from a greater diversity of collections before publishing subsequent volumes of this series. While not without its flaws, I think this is an attractive, useful, and generally well- executed book. As stated at the outset, it is a clear improvement over D’Abrera’s Butterflies of the Neotropical Region, at least for Venezuela, and will be a helpful tool for curating even large butterfly collections like those of the AMNH and NMNH. Given the complexity of the neotropical butterfly fauna, and the extent of our ig- norance regarding its systematics and biogeography, Neild is to be congratulated for attempting to match DeVries’ (1987) standard with a guide to a larger and more diverse South American country. — Andrew V. Z. Brower, Dept, of Entomology, Na- tional Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. LITERATURE CITED Cracraft, J. 1983. Species concepts and speciation analysis. Current Ornithol. 1:159-187. DeVries, P. J. 1987. The butterflies of Costa Rica and their natural history. Papilionidae, Pieridae, Nymphalidae. Princeton University Press, Princeton, N.J. Jenkins, D. W. 1990. Neotropical Nymphalidae VIII. Revision of Eunica. Bull. Allyn. Mus. 131:1-177. de Jong, R., Vane- Wright, R. I., and P. R. Ackery. 1996. The higher classification of butterflies (Lepidoptera): problems and prospects. Ent. Scand. 27:65-101. Lamas, G. 1996. Diez notas sinonimicas sobre Satyrinae neotropicales, con la descripcion de 1996 BOOK REVIEWS 239 dos subespecies nuevas de Peru y Ecuador (Lepidoptera; Nymphalidae). Rev. Per. Ent. 39:49-54. Martin, J. A., and D. P. Pashley. 1992. Molecular systematic analysis of butterfly family and some subfamily relationships (Lepidoptera: Papilionoidea). Ann. Ent. Soc. Am. 85:127- 139. Mayr, E. 1940. Speciation phenomena in birds. Am. Nat. 74:249-278. Nixon, K. C. and Q. D. Wheeler. 1990. An amplification of the phylogenetic species concept. Cladistics 6:211-223. Otero, L. D. 1990. Estudio de algunos caracteres para su uso en la clasificacion de Eurytelinae (Lepidoptera: Nymphalidae). Bol. Entomol. Venezol. N. 8.5:123-138. 1996 HONORARY MEMBERS Dr. F. S. Chew Ms. Su Zan Noguchi Swain Dr. James Forbes Dr. James A. Slater Dr. John G. Franclemont LIFE MEMBERS Dr. Annette Aiello Professor Clive R. Bailey Mr. David G. Casdorph Mr. Franklyn W. Commisso Dr. Howard E. Evans Dr. Durland Eish Mr. Irving Granek Prof. Dipl. Ing. Ernst Heiss Dr. Donald F. J. Hilton Professor Hussein S. Hussein Mr. Mark Indenbaum Dr. Kurt Johnson Dr. Gary G. Kennen Dr. J. Krikken Mr. George Ladd Dr. Brobson Lutz, M.D. Dr. Gordon A. Marsh Dr. John C. 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Society members will be charged a fee of $23.00 per printed page and $8.50 per plate of figures. Non-members will be charged $65.00 per printed page and $15.00 per plate of figures. Member authors who are students, or who do not have institutional affiliation may petition to the Society for waiver of page charges for no more than eight pages on a once a year basis. Because of limited funds, all such requests will be handled on a first-come first-serve basis. Authors will receive a reprint order blank with the proofs. Reprints are ordered directly from the printer with no benefit accruing to the Society. Journal of the New York. Entomological Society VOLUME 104 SUMMER-AUTUMN 1996 NOS. 3-4 CONTENTS Systematics of the New World waterscorpion genus Curicta Stal (Heteroptera: Nepidae) Steven L. Kejfer 117-215 A new neotropical fungus gnat (Diptera: Sciaroidea: Keroplatidae) with myrme- cophagous larvae Loi'c Matile 216-220 Two new species of Megastylus from the New World (Hymenoptera: Ichneumon- idae; Orthocentrinae) David B. Wahl 221-225 Notes and Comments Myrmecophily in Keroplatidae (Diptera: Sciaroidea) Annette Aiello and Pierre Jolivet 226-230 Book Reviews Studies on Hemipteran Phylogeny Randall T. Schuh 231-235 The Butterflies of Venezuela, Part 1: Nymphalidae I (Limenitidinae, Apaturinae, Charaxinae) Andrew V. Z. Brower 236-239 Honorary, Life, and Sustaining Members 240 Reviewers for 1996 240 WINTER-SPRING 1997 Nos. 1-2 Journal of the New York Entomological Society (ISSN 0028-7199) Vol. 105 Ql HQ Devoted to Entomology in General JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Editor; James M. Carpenter, Department of Entomology, American Mu- seum'* of Natural History, Central Park West at 79th Street, New York, NY 10024 Book Review Editor; James S. 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Soc. 105( 1-2): 1-14, 1997 AN ANALYSIS OF THE GENUS STAL WITH DESCRIPTION OF FOUR SPECIES, AND SOME TAXONOMIC REARRANGEMENTS (HEMIPTERA: HETEROPTERA; LARGIDAE) * ' Harry Brailovsky and Cristina Mayorga Institute de Biologia, UNAM, Depto. de Zoologia, Apdo Postal No. 70-153, Mexico 04510, D.E, Mexico Abstract. — Four new species of Stenomacra Stal from Costa Rica, Ecuador, and Brazil are described and illustrated. Stenomacra cliens (Stal), Stenomacra gracilis Schmidt, and Steno- macra marginella van mexicana Schmidt are synonymized under Stenomacra marginella (Her- rich-Schaeffer); Theraneis ferruginea Mayr is transferred to the genus Stenomacra and syn- onymized under Stenomacra scapha (Perty); Theraneis dissimilis Distant is transferred to the genus Stenomacra, resulting on the new combination Stenomacra dissimilis (Distant). New distributional data are included for each species. A key to the known species of Stenomacra is also given. Key words: Insecta, Heteroptera, Largidae, Stenomacra, new species, Costa Rica, Ecuador, Brazil. The Neotropical genus Stenomacra Stal (1870) had never been revised. However, the discovery of four undescribed species and several taxonomical problems made necessary the present analysis, which incorporates new distnbution data and a key to separate the known species. The genus is characterized by a slender to elongate body, nearly parallel-sided, with the head in dorsal view and between eyes flat or slightly convex, eyes small and barely pedunculate, ocelli absent, antennal segment I uniformly slender, bucculae short, slightly elevated, and scarcely extended backward as far as the antenniferous tubercle, rostrum just reaching the posterior margin of the mesothorax, pronotum with the anterior lobe not globose and the humeral angles rounded, hemelytra ma- cropterous or eventually submacropterous, anterior coxae unarmed and fore femora armed with two or three subdistal spines (middle and hind femora unarmed), me- tathoracic peritreme never auriculate, and the abdominal sternite VII of the female mesally cleft. Largulus Hussey (1927) is somewhat similar to Stenomacra, but the rostrum reaches the metathorax, and the body and legs are remarkably more slender. Previously, only five species of Stenomacra, S. cliens (Stal), S. gracilis Schmidt, S. limbatipennis (Stal), S. marginella (Herrich-Schaeffer) and S. scapha (Perty), and one variety, S. marginella mexicana Schmidt were known (Hussey and Sherman, 1929 and Schmidt 1931). In this contribution we add four new species collected in Costa Rica, Brazil and Ecuador; one species, Theraneis dissimilis Distant (1883), is transferred to the genus Stenomacra, forming the new combination Stenomacra dis- similis', Theraneis ferruginea Mayr (1865) is also transferred to Stenomacra but '2 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) y synonymized under S. scapha; and Stenomacra cliens, S. gracilis and S. marginella var. mexicana are placed as new synonyms within S. marginella. After this contribution, the current number of species belonging to Stenomacra is fixed at eight. The following institutional abbreviations are used in the text: American Museum of Natural History, New York (AMNH); the Natural History Museum, England (BMNH); California Academy of Sciences, San Francisco, California (CAS); Insti- tuto Nacional de Biodiversidad, Santo Domingo de Heredia, Heredia, Costa Rica (INBIO); Museo Miguel Lillo, Tucuman, Argentina (MLTA); Museum National d’Histoire Naturelle, Paris, France (MNP); Museum National, Rio de Janeiro, Brazil (MNR); Naturhistorisches Museum, Wien, Austria (NMW); Naturhistoriska Riks- museet, Stockholm, Sweden (NRE); Texas A. & M. University, College Station (TAMU); Instituto de Biologia, Universidad Nacional Autonoma de Mexico (UNAM); Utah State University, Utah (USU). Synonyms and references of Stenomacra should be consulted in the catalogs of Hussey and Sherman (1929) and Henry and Froeschner (1988) to avoid repetition and save space. All measurements are in millimeters. Stenomacra turrialbana, new species Fig. 1 Female. Dorsal coloration. Head bright orange brown, with tylus and antennal seg- ments I to IV black; pronotum light orange brown with sides of apical one third yellowish, center longitudinally and collar light orange brown, and basal third me- dially yellowish; scutellum light orange brown with a wide yellow longitudinal stripe extending from the base to apex; hemelytra light orange brown with following areas yellow: costal margin, basal angle of the corium, inner angle of the apical margin of corium, claval commissure, and anal border; hemelytral mxmbrane ambarine, with outer margin darker; connexival segments with upper margin yellow, and lower one orange brown; abdominal segments bright orange brown. Ventral coloration. Bright reddish brown to orange brown, with bucculae, mesial gular region, prosternum, mesostemum and metasternum, upper third of acetabulae, anterior margin of me- sothorax and metathorax, close stripe near posterior margin of mesothorax and meta- thorax, and the area neighbouring to metathoracic peritreme black; following areas yellow: quadrate spot in the anterior margin of prothorax, metathoracic peritreme, mesial and wide longitudinal stripe running from III to VI abdominal sterna, pleural margin of abdominal sterna III to VII, paratergite VIII and external margin of par- atergite IX; rostral segments I and IV bright black, II bright orange brown, and III bright orange brown with apical third bright black; legs reddish black, with trochan- ters dirty yellow. Structure. Body slender, nearly parallel- sided; head, pronotum, scutellum, clavus, corium, thorax, and abdominal sterna with long, slender, erect setae, intermixed with short decumbent silvery setae. Head. Dorsally between eyes almost flat; eyes barely pedunculate; bucculae short, elevated, extending only to anterior margin of antenniferous tubercles; rostrum reaching posterior margin of mesothorax. Thorax. Pronotum. Trapezoidal, wider than long, and markedly bilobed with transverse impression deeply punctate; collar wide; anterior pronotal lobe with lateral margins rounded; posterior pronotal lobe with lateral margins obliquely 1997 THE GENUS STENOMACRA 3 Fig. 1. Stenomacra turrialbana, new species, dorsal view. straight; anterior lobe impunctate; posterior lobe with scattered punctation; posterior border straight; humeral angles rounded. Legs. Covered with fine adpressed silvery setae, intermixed with long silvery setae; fore femora armed with three single ranked, of long subapical spines; middle and hind femora unarmed. Scutellum. Triangular, slightly longer than wide, and flat; each side of anterior third of scutellar disc punc- tate. Hemelytra. Macrpterous, reaching the apex of the last abdominal segment; cla- vus and corium punctate; costal margins virtually straight. Measurements. Length head: 2.00; width across eyes: 2.12; interocular space: 1.20; preocular distance: 1.08; length antennal segments: I, 3.72; II, 2.12; III, 1.32; IV, 2.68. Pronotal length: 2.68; maximum width of anterior lobe: 2.40; maximum width of posterior lobe: 3.76. Scutellar length: 1.72; width: 1.66. Total body length: 14.10. Variation. 1. Rostral segments II and III bright black. 4 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Male. Unknown. Holotype female. COSTA RICA: PROVINCIA CARTAGO: Grano de Oro, Chir- ripo, Turrialba (1120 mts.), IX.1992. P. Campos. Deposited in INBIO. Paratypes. Five females: COSTA RICA: PROVINCIA CARTAGO: Grano de Oro, Chirripo, Turrialba (1120 mts.), 8-30. VII. 1992, 8-31. VIII. 1992 and IX.1992. P. Cam- pos. Deposited INBIO and UNAM. Discussion. This new species is very distinct. The pronotum is light orange brown with the sides of its apical third, as well as a wide longitudinal stripe extending from its base to the middle third, yellowish. Stenomacra dissimilis (Distant), also recorded from Costa Rica, has the corium black with costal and apical margins, as well as the basal angle and claval suture yellow. The pronotal disc has another type of coloration. In S. turrialbana the corium is light orange brown with only the costal margin, the basal angle and the inner angle of the apical margin yellow. Etymology. Named for the type locality, Turrialba. Stenomacra tungurahuana, new species Fig. 2 Male. Dorsal coloration. Head, including antennal segments and rostral segments black, with basal join of antennal segment I red orange; pronotum red orange with callar region black; scutellum black with apex red orange; clavus and corium red orange; hemelytral membrane white; connexival segments black with upper border red orange; abdominal segments black. Ventral coloration. Black, with the external margin of bucculae, anterior, lateral and posterior margin of prothorax, apex of me- tathoracic peritreme, pleural margins of abdominal sterna and posterior margin of the genital capsule red orange. Structure. Similar to S. turrialbana. Rostrum reaching anterior third of metathorax. Scutellar disc scattered punctate. Genital capsule. Pos- teroventral edge entire, almost straight. Measurements. Length head: 1.44; width across eyes: 1.84; interocular space: 1.08; preocular distance: 1.03; length antennal segments: I, 3.28; II, 1.88; III, 1.12; IV, 2.76. Pronotal length: 2.00; maximum with of anterior lobe: 2.08; maximum width of posterior lobe: 3.44. Scutellar length: 1.60; width: 1.56. Total body length: 12.85. Female. Coloration. Similar to the male. Connexival segments VIII and IX red orange; abdominal segment VIII black with posterior margin red orange, and IX red orange; genital plates black with external margin red orange. Measurements. Length head: 1.56; width across eyes: 1.88; interocular space: 1.20; preocular distance: 1.00; length antennal segments: I, 2.96; II, 1.80; III, 1.08; IV, 2.40. Pronotal length: 2.28; maximum width of anterior lobe: 2.12; maximum width of posterior lobe: 3.32. Scutellar length: 1.40; width: 1.52. Total body length: 12.90. Variation. 1. Posterior margin of abdominal stemite VII red orange. Holotype male. ECUADOR: PROVINCIA TUNGURAHUA: Banos, 24.VII.75. J. Longino. Deposited in UNAM. Paratypes. Two males, two females: ECUADOR: PROVINCIA TUNGURAHUA: Banos (1800 mts.), 4-5.IV.58. W. Weyrauch. Deposited in MLTA and UNAM. One female: ECUADOR: Banos, 20.11.37. S. W. Frost. Deposited in USNM. One male, one female: ECUADOR: Quito, 1930. R. Benoist. Deposited MNP. Discussion. As in Stenomacra scapha (Perty), the clavus and corium are red orange. 1997 THE GENUS STENOMACRA 5 Fig. 2. Stenomacra tungurahuana, new, species, dorsal view. and the hemelytral membrane is white. However S. tungurahuana is macropterous, the hemelytra reaching the apex of the last abdominal segment; antennal segments I to III, the callar region of the pronotal disc, and the tibiae are black. S. scapha is submacropterous, the hemelytra reaching the posterior margin of abdominal segment VI; antennal segments I to III, the callar region and the tibiae are red orange to pale orange. Etymology. Named for its occurrence on Tungurahua. Stenomacra magna, new species Fig. 3 Male. Dorsal coloration. Head including antennal segment I to III bright orange chestnut; antennal segment IV with anterior half bright orange chestnut, and posterior half dark brown; pronotum light yellow with callar region and punctures bright orange chestnut; scutellum bright orange chestnut; clavus and corium light yellow with diffuse brown shadow area; hemelytral membrane light ambarine; connexival segments bright orange with upper and lower margin yellow; abdominal segments 6 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Fig. 3. Stenomacra magna, new species, dorsal view. bright orange. Ventral coloration. Head including rostral segments I to III bright orange chestnut; rostral segment IV black with basal third bright orange chestnut; thorax bright orange chestnut with sternal region black, and following areas yellow: acetabulae, anterior and posterior margin of the prothorax, posterior margin of ma- sothorax and metathorax, and metathoracic peritreme; legs bright orange; abdominal sterna bright orange, with pleural margin yellow, and anterior margin of each stemite black; genital capsule bright orange, with yellow spot on the middle of the body. Structure. Similar to S. turrialbana. Rostrum reaching posterior third of mesothorax; scutellum wider than long or as longer as wide. Genital capsule. Posteroventral edge entire, almost straight. Measurements. Length head: 1.40; width across eyes: 2.00; interocular space: 1.16; preocular distance: LOO; length antennal segments: I, 3.44; II, 1.64; III, 1.06; IV, 1997 THE GENUS STENOMACRA 1 2.80. Pronotal length: 2.24; maximum width of anterior lobe: 2.08; maximum width of posterior lobe: 3.08. Scutellar length: 1.32; width: 1.32. Total body length: 13.00. Female. Color. Similar to male. Connexival segments, abdominal segments, and genital plates bright orange yellow; metathoracic peritreme bright orange. Measure- ments. Length head: 1.48; width across eyes: 2.08; interocular space; 1.20; preocular distance: 1.00; length antennal segments: I, 3.56; II, 1.80; III, 1.08; IV, 2.64. Pronotal length: 2.44; maximum width of anterior lobe: 2.24; maximum width of posterior lobe: 3.56. Scutellar length: 1.48; width: 1.56. Total body length: 14.40. Holotype male. BRAZIL: NOVA TEUTONIA: Santa Catarina, 27 lLN-52 23'W, XII. 1967. F. Plaumann. Deposited in TAMU. Paratype female. One female: BRAZIL: NOVA TEUTONIA: Santa Catarina, 27 ll'N- 52 23'W, 28.1.1967. E Plaumann. Deposited in UNAM. Discussion. This unique species can be distinguished by having antennal segments I to III bright orange, and IV bicolorous, the clavus and corium light yellow with a diffuse brown area, and the legs entirely bright orange. In Stenomacra limbatipennis (Stal), antennal segments I to IV and the legs are black to reddish brown, and the corium is never entirely yellow; the clavus of S. limbatipennis is black with the claval commissure yellow. Etymology. Named for its large size; from the Latin word, magnus. Stenomacra atra, new species Fig. 4 Male. Dorsal coloration. Head, antennal segment I, anterior pronotal lobe, and scu- tellum (apex dark orange) black; antennal segments II to IV yellow with basal third of II and apical third of IV reddish brown to black; posterior pronotal lobe dark orange red; clavus yellow; corium yellow with the apical margin and a longitudinal median stripe, covered the endocorium reddish brown; hemelytral membrane dirty white; connexival segments I to V orange, and VI and VII black, with upper and anterior margin orange; abdominal segments orange with diffuse brownish spots. Ventral coloration. Including rostral segments I to IV and legs reddish brown to black; posterior margin of prothorax orange brown; tarsi bright chestnut orange; abdominal sterna with pleural margins III to VII, and posterior margin of VII orange; metathoracic peritreme with the lobe dirty yellow; genital capsule black with median body yellow. Structure. Similar to S. turrialbana. Rostrum reaching anterior third of metathorax. Scutellar disc scattered punctate. Genital capsule. Posteroventral edge entire, narrow, and almost straight. Pubescence. Head, exposed parts of thorax, ab- domen, scutellum, legs, clavus and corium with short, fine, suberect pubescence, intermixed with long erect hairs; head in dorsal view densely set with silvery ad- pressed hairs; thoracic pleura and lateral areas of abdominal stemite III to V, and eventually VI with patches of very dense, felted silvery pubescence. Measurements. Length head: 1.60; width across eyes: 2.16; interocular space: 1.20; preocular distance: 1.12; length antennal segments: I, 3.80; II, 1.84; III, 1.18; IV, 3.20. Pronotal length: 2.40; maximum width of anterior lobe: 2.20; maximum width of posterior lobe: 3.20. Scutellar length: 1.44; width: 1.32. Total body length: 14.10. Female. Color and structure similar to the male. Anterior pronotal lobe black with callar region bright orange; scutellum dark orange with lateral margins black; ab- 8 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Eig. 4. Stenomacra atra, new species, dorsal view. dominal segments light orange with anterior margin brown; abdominal sterna reddish brown with pleural margins III to V yellow, and VI and VII dark orange to reddish brown; genital plates yellow. Measurements. Length head: 1.68; width across eyes: 2.24; interocular space: 1.34; preocular distance: 1.12; length antennal segments: I, 3.92; II to IV absent. Pronotal length: 2.56; maximum width of anterior lobe: 2.42; maximum width of posterior lobe: 3.40. Scutellar length: 1.48; width: 1.42. Total body length: 14.52. Holotype male. BRAZIL: Bahia (without data). Deposited NMW. Paratype. One female: BRAZIL (without data). Deposited UNAM. Discussion. This new species superhcially resembles Stenomacra limbatipennis (Stal) in size and shape. Both have completely reddish brown to black legs. It is however readily distinguishable by the yellow clavus, as well as by the yellow corium with its apical margin and a large irregular elongate brown stripe. In addition 1997 THE GENUS STENOMACRA 9 the hemelytral membrane is dirty white. In S. limbatipennis the clavus and corium are black with the anal border, claval commissure, and costal margin of the corium yellow, with the hemelytral membrane dark brown. Etymology. From the Latin, atra, black. Stenomacra limbatipennis (Stal) Theraneis limbatipennis Stal, 1860: 45 This is a readily recognizable species by virtue of its completely black to reddish brown antennal segments I to IV, and legs, and by having the pronotum, scutellum, and hemelytra black with following areas yellow: posterior margin of pronotum, apex of scutellum, anal border and claval commissure, and costal margin of the corium; hemelytral membrane dark brown. The relation of this species with S. atra is given under the preceeding species. Types. Stal (1860) described this species from a single female specimen. The type of Theraneis limbatipennis, housed in the Naturhistoriska Riksmuseet, Stockholm, Sweden, was examined. The label data for the holotype specimen is Brazil, F. Sahib. The type locality is Rio de Janeiro. Bergroth (1905) recorded this species from Minas Gerais, Brazil. Material examined. One male from Brazil (without data) and one female from, BRAZIL. RIO DE JANEIRO, Itatiaya Maromba, 20.1. 1925. J. F. Zikan. Deposited in AMNH and UNAM. Distribution. Brazil. Stenomacra dissimilis (Distant), New Combination Theraneis dissimilis Distant, 1883: 225. Theraneis dissimilis Brailovsky, 1991: 636. Individuals of this species are medium sized with the head (bucculae black or yellow), antennal segments I to IV, and legs black; pronotal disc black with each margin yellow; corium yellow with a large, irregular and elongate median black spot; hemelytral membrane dark brown, and connexival segments light yellow. Stenomacra marginella (Herrich-Schaeffer) has antennal segments I to IV, tibiae and tarsi black, but the antenniferous tubercles, coxae, trochanters, and femora (ex- cept the black dorsal subapical face) light orange yellow, and the connexival seg- ments are black with upper margin yellow. Types. Distant (1883) described Theraneis dissimilis from at least three specimens, two from Costa Rica (Cache) and one from Panama (Bugaba). The types are housed in the Natural History Museum, London. A female paralectotype was examined. The label data are Panama, Bugaba, Champion. Material examined. One female, COSTA RICA: Provincia Alajuela, sector San Ramon, 620 mts., 1 1-I5.IV.I994. E. Lopez. Deposited in INBIO. Two females, COS- TA RICA: Provincia Alajuela, 20 km., S. Upala, 1-10. VIII. 1991 and 1 0-2 l.V. 1991. F. D. Parker. Deposited in USU. Six females, COSTA RICA, Provincia Guanacaste, Parque Nacional Guanacaste, Estacion Pitilla, 9 km., S. of Sta. Cecilia, 700 mts., VII.1988, 1.1989, XII.1989, 11.1990, 29.IV.1992. Deposited in INBIO and UNAM. One female, COSTA RICA, Provincia Guanacaste, Estacion Cacao, SW., side of 10 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Volcan Cacao, 1000-1400 mts., (Malaise Trap), 1988-1989 (without data). Deposited in INBIO. Distribution. Costa Rica and Panama. Stenomacra marginella (Herrich-Schaeffer) Largus marginellus Herrich-Schaeffer, 1850: 182. Theraneis aliens Stal, 1862: 315. NEW SYNONYMY. Stenomacra marginella: Stal, 1870: 98. Stenomacra aliens: Stal, 1870: 98. Stenomacra aliens: Hussey & Sherman, 1929: 25. Stenomacra marginella: Hussey & Sherman, 1929: 26. Stenomacra marginella: Schmidt, 1931: 44. Stenomacra marginella var. mexicana Schmidt, 1931: 44. New Synonymy. Stenomacra gracilis Schmidt, 1931: 45. New Synonymy. Stenomacra marginella: Blote, 1931: 99. This is a readily recognizable species by virtue of having antennal segments I to IV, rostral segments I to IV, tibiae and tarsi completely black to reddish brown, and by having the following areas always orange to orange yellow: antenniferous tuber- cles, bucculae, pronotal margins including the collar, costal margin of the corium, anal suture and commissure claval, apex of scutellum, upper margin of connexival segments, acetabulae, coxae, and trochanters. Variation. This species is extremely variable in coloration, and several color forms can be found into the same population. 1. Head in dorsal view black or light orange, or only with the space close to the eyes black. 2. Anterior lobe of the pronotum orange or yellow and posterior lobe black, with or without median longitudinal orange to yellow stripe, or anterior and posterior lobe black with a median longi- tudinal yellow stripe, or both lobes with a diffuse yellow mark on a light brown background. 3. Scutellum black with the apex orange to yellow, or scutellar disc orange to yellow with basal third black. 4. Femora completely black or reddish brown, or completely bright orange, or well intermixed with basal third orange and the rest black, or orange with only the dorsal face of the apical third black, and other variants. Types. Herrich-Schaeffer (1850) described Largus marginellus from Brazil but the type could not be located. The description, however, is adequate enough to fit this species. Stal (1862) described Theraneis aliens from Mexico and the type located in the Naturhistoriska Riksmuseet, Stockholm, was examined. Schmidt (1931) de- scribed Stenomacra marginella var. mexicana from Mexico and Stenomacra gracilis from Colombia but the types could not be located, however more than 150 specimens were examined and compared with the original descriptions which fit this species. Until further evidence is discovered, it seems best to produce these nominal taxa as synonyms due to the lack of morphological characters that would allow us to separate the taxa, as well as the extraordinary chromatic variation found among the long series of specimens examined. Walker (1873) recorded this species from two Mexican localities Orizaba in the State of Veracruz and from Oaxaca. Distant (1880-1893) recorded this species from Mexico including Coahuila (Parras, and Saltillo); Jalisco; Morelos (Cuernavaca); 1997 THE GENUS STENOMACRA 11 Guerrero (Amula, Chilpancingo, and Xucumanatlan); Veracruz (Jalapa, and Orizaba). Guatemala (Capetillo, and San Geronimo). Nicaragua (Chontales). Panama (Volcan de Chiriqui). Colombia (Bogota, and Antioquia). As well as from Brazil. Barber (1910) mentioned this species from Arizona (Huachuca Mts.) in the United States of America, and Bldte (1931) from California (United States of America) and from San Jose (Costa Rica). Henry and Froeschner (1988) in their catalogue recorded this species from the United States of America (California, Arizona, and New Mexico). Material examined. Three males from UNITED STATES OF AMERICA: Arizona, Portal (S.W. Res. Sta.) (5400'), 10. VI. 1969. W. M. Nunes and T. H. Halstead. De- posited UNAM. Two males, three females from MEXICO: Nayarit, km., 98 road Guadalajara-Tepic, 27. VII. 1984. E. Barrera and A. Ibarra. Deposited UNAM. One male, three females from MEXICO: Durango, 10 mi., N.W. Las Nieves (6500'), 1. VII. 1971. C. W. O’Brien. Deposited UNAM. One male, one female from MEXI- CO: Durango, Presa Pena, 6. VIII. 1979. E. Barrera. Deposited UNAM. One male, one female from MEXICO: Colima, Colima, IX. 1940. C. Hoffmann. Deposited UNAM. One male from MEXICO: Jalisco, Teocatliche, 22.VIII.1970. B. Villegas. Deposited UNAM. Three males from MEXICO: Jalisco, km., 93 road Guadalajara- Ixtlan, 7. VII. 1982. H. Brailovsky. Deposited UNAM. Five females from MEXICO: Jalisco, Chamela, 3.XII.1976. H. Brailovsky. Deposited UNAM. One male, two fe- males from MEXICO: Michoacan, 8 mi., W. of Patzcuaro (7600'), 14. VIII. 1982. O’Brien and Wibmer. Deposited UNAM. One female from MEXICO: Guerrero, Chapa, 26. VII. 1975. R. Medellin. Deposited UNAM. One female from MEXICO: Guerrero, Chichihualco (1250 mts.), 6. VII. 1979. H. Brailovsky. Deposited UNAM. 19 males, 14 females from MEXICO: Morelos, Cuautla, 17.VI.1966, 22.VI.1969, 8- 12.V.1970, 28.VIII.1970, 15.VIII.1984, 20.V.1989. H. Brailovsky. Deposited UNAM. Six males, 8 females from MEXICO: Morelos, Yautepec, 6. VI. 1981. H. Brailovsky. Deposited UNAM. Nine males, eleven females from MEXICO: Morelos, Tepoztlan, 6.VI.1976, 25.11.1977, 9.IX.1977, 5.V.1980, 3.IV.1981. E. Marino, E. Gonzalez, J. Butze, E. Barrera. Deposited UNAM. Two females from MEXICO: Morelos, Tetela del Volcan, 30. VI. 1982. A. Ibarra. Deposited UNAM. Two males, one female from MEXICO: Estado de Mexico, Malinalco, 28. IV. 1984. H. Brailovsky. Deposited UNAM. Two males, four females from MEXICO: Estado de Mexico, Valle de Bravo, 3.VII.1969, 18.V.1973. H. Brailovsky. Deposited UNAM. 12 males, 17 females from MEXICO: Distrito Federal, Pedregal de San Angel (Jardin Botanico, UNAM), 17.IV.1968, 11.V.1968, 12.VI.1968. H. Gonzalez Almada. Deposited UNAM. One male, one female from MEXICO: Distrito Federal, Jalpan, 26. IV. 1975. C. Beutel- spacher. Deposited UNAM. Three males, three females from MEXICO: Distrito Fed- eral, Xochimilco (Tepepan), 10.V.1969, 11.IV.1975, 2.VII.1990. L. Rivera, W. Brai- lovsky, E. Martin. Deposited UNAM. Two males, three females from MEXICO: Hidalgo, Tasquillo, 15. VII. 1979. H. Brailovsky. Deposited UNAM. Two males from MEXICO: Hidalgo, Tecozautla, 18.V.1996. E. Barrera, H. Brailovsky. Deposited UNAM. Three females from MEXICO: Hidalgo, Santiago de Anaya, 21. IV. 1990. C. Mayorga. Deposited UNAM. Two females from MEXICO: Guanajuato, Guanajuato, 29. V. 1975. P. Rojas. Deposited UNAM. One female from MEXICO: Guanajuato, Salamanca, 25.IX.1975. J. Bueno. Deposited UNAM. Two males from MEXICO. Queretaro, Queretaro, 6.IV.1977. J. Garcia Figueroa. Deposited UNAM. Three males, one female from MEXICO: Puebla, Tecamachalco, 27.11.1992. G. Ortega Leon, C. 12 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Mayorga, E. Barrera. Deposited UNAM. Four females from MEXICO: Puebla, San Esteban (Necoxcalco), 27. IX. 1992, 15. X. 1992. E. Barrera, H. Brailovsky, C. May- orga. Deposited UNAM. Two males, three females from MEXICO: Puebla, Tehu- acan, 27.11.1992, 27. IX. 1992. E. Barrera, C. Mayorga, G. Ortega Leon. Deposited UNAM. One female from MEXICO: Puebla, Atlixco (La Trinidad), 25.IV. 1994. E. Barrera, H. Brailovsky. Deposited UNAM. Three males, four females from MEXI- CO: Veracruz, Las Minas, 16. IX. 1982. E. Barrera, H. Brailovsky. Deposited UNAM. One male, one female from MEXICO: Veracruz, Estacion de Microondas de Fortin, 15.VII.1980. E. Barrera. Deposited UNAM. One male from MEXICO: Veracruz, Conejos, 13. VIII. 1984. H. Brailovsky. Deposited UNAM. One female from MEXI- CO: Veracruz, Orizaba (1880 mts.), 15. VIII. 1975. A. N. Garcia Aldrete. Deposited UNAM. One female from MEXICO. Oaxaca, Mitla, 26.V.1970. H. Brailovsky. De- posited UNAM. One male from MEXICO: Oaxaca, Portillo del Rayo (Jalatengo), 21.x. 1982. H. Brailovsky. Deposited UNAM. One male, one female from MEXICO: Oaxaca, Monte Alban, 27. III. 1990. E. Barrera, A. Cadena. Deposited UNAM. One female from MEXICO: Oaxaca, km., 193 road Teotitlan-Oaxaca, 15. III. 1989. A. Cadena, L. Cervantes. Deposited UNAM. Three males, seven females from MEX- ICO: Oaxaca, km., 47 road Huajuapan de Leon-Oaxaca (2010 mts.), 8.VII.1995. E. Barrera, G. Ortega Leon. Deposited UNAM. One male from MEXICO: Chiapas, Union Juarez, 21. IV. 1983. E. Barrera. Deposited UNAM. One female from MEXI- CO: Chiapas, Reserva El Triunfo, 6. VII. 1993. C. Mayorga. Deposited UNAM. Three males, three females from COSTA RICA: San Jose (Sta. Maria de Dota), 30.IV. 1984. E. Barrera. Deposited UNAM. Two females from COLOMBIA: 27 mi.. South of Pasto Narino (1960 mts.), 28.11.1955. E. I. Schlinger, E. S. Ross. Deposited CAS. One female from VENEZUELA: Merida, km., 20 road Merida-El Morro, 20.XI.1992. Deposited UNAM. Stenomacra scapha (Perty) Lygaeus scapha Perty, 1833: 172. Largus incisus Herrich-Schaeffer, 1850: 182. Theraneis ferruginea Mayr, 1865: 436. New Synonymy. Brailovsky (1991) revised the genus Theraneis Spinola. He described four new species and included a key to separate the known taxa. However, T. constricta Stal, T. ferruginea Mayr, and T. isobel Hussey were not included in the key since there were no specimens available and the corresponding types not located at the time. Recently, the authors had the opportunity to examine the type of T. ferruginea, which is deposited in the Naturhistorisches Museum, Wien. As a result, this species has been transferred to the genus Stenomacra and synonymized under S. scapha (Perty). The latter species is distinguished by submacropterous individuals with whitish he- melytral membrane; also the head, antennal segments, pronotum, legs and abdomen are reddish brown. Its distributional range is restricted to Brazil, the same area from where its new synonym was collected. Types. Perty (1833) described this species from a single male specimen. The type was not located, but the description and figure, are adequate to determine its identify. The type specimen of iMrgus incisus Herrich-Schaeffer (1850) was not examined, but the figure in the original description is of a specimen identical to S. scapha. 1997 THE GENUS STENOMACRA 13 Mayr (1865) described Theraneis ferruginea and the type specimen is a female located in the Naturhistorisches Museum, Wien. The type was examined and it has the following label data: Brasilien, Schott. The latter species has been transferred to the genus Stenomacra and synonymyzed under S. scapha. Hussey (1929) recorded this species from Rio de Janeiro and Minas Geraes in Brazil. Material examined. One female from, BRAZIL: Petropolis, II. 1857. In MNP. One male, two females from BRAZIL: Rio de Janeiro, Itatiaya, 1200 mts., 26.XII.1953, Seabra and Alvarenga. Deposited in AMNH and UNAM. Three males, one female from BRAZIL: Rio de Janeiro, Itatiaya, 29. XII. 1950. C. Leite, Seabra and Zickan. Deposited in MNR and UNAM. KEY TO STENOMACRA SPECIES 1. Antennal segment I bright orange 2 1' Antennal segment I black to dark reddish brown 3 2. Antennal segment IV black with basal third orange; submacropterous, with the he- melytral membrane reaching anterior margin of abdominal segment VI; posterior lobe of the pronotum, clavus and corium light orange red (Brazil) scapha (Perty) 2' Antennal segment IV orange with apical third black; macropterous, with the hemelytral membrane almost reaching the apex of the last abdominal segment; posterior lobe of the pronotum, clavus and corium yellow (Brazil) magna, new species 3. Antennal segments II and III yellow (Brazil) atra, new species 3' Antennal segments II and III black 4 4. Clavus and corium bright orange red; hemelytral membrane white (Ecuador) tungurahuana, new species 4' Clavus and corium never bright orange red; hemelytral membrane light brown 5 5. Anterior margin of the pronotum and collar black to bright reddish brown 6 5' Anterior margin of the pronotum and collar yellow to orange 7 6. Pronotum black with humeral angles and posterior margin yellow; scutellum black with apex ochre; lower margin of acetabulae yellow (Brazil) limbatipennis (Stal) 6' Pronotum light orange brown with sides of apical one third, and basal third medially yellowish; humeral angles light orange brown; scutellum light orange brown with a wide yellow longitudinal stripe running from the base to the apex; lower margin of acetabulae reddish brown (Costa Rica) turrialbana, new species 7. Coxae and trochanters black; antenniferous tubercle black (Costa Rica, Panama) . . . dissimilis (Distant) 7' Coxae and trochanters bright orange yellow; antenniferous tubercle bright orange yel- low (United States of North America, Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Brazil) marginella (Herrich-Schaeffer) ACKNOWLEDGMENTS We would like to thank the following individuals and institutions for their help in lending specimens pertinent to this project: Dr. R. T. Schuh (AMNH); Mrs. Janet Margerison-Knight (BMNH); Dr. Norman D. Penny (CAS); Mr. Jesus Armando Ugalde Gomez (INBIO); Dra. Maria del Valle Ajmat de Toledo (MLTA); Dra. Dominique Pluot (MNP); Dr. Jose Candido de Melho Carvalho (MNR); Dra. Ulrike Aspock and Dr. Herbert Zettel (NMW); Dr. Per Lindskog (NRE); Dr. Joseph C. Schaffner (TAMU); Dr. Wilford J. Hanson (USU). Special thanks to Dr. Manuel Balcazar (UNAM), Dr. Atilano Contreras (UNAM), and Biol. Ernesto Barrera (UNAM) for their comments on an early version of the manuscript, and to Mr. Albino Luna for the preparation of the dorsal view illustration of Stenomacra turrialbana. 14 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) LITERATURE CITED Barber, H. G. 1910. Some Mexican Hemiptera-Heteroptera new to the fauna of the United States. J. N. Y. Entomol. Soc. 18:34-39. Bergroth, E. 1905. Rhynchota Neotropica. Rev. d’ Ent. 24:104-112. Blote, H. C. 1931. Catalogue of the Pyrrhocoridae in ’S Rijks Museum van Natuurlijke His- toric. Zool. Meded. 14:97-136. Brailovsky, H. 1991. Four new species of the Neotropical genus Theraneis Spinola (Hemiptera: Heteroptera: Largidae). J. New York Entomol. Soc. 99(4):630-636. Distant, W. L. 1880-1893. Insecta. Rhynchota. Hemiptera-Heteroptera. Biologia Centrali- Americana. London: 1-462. Henry, T. J., and R. C. Froeschner. 1988. Catalog of the Heteroptera or true bugs, of Canada and the Continental United States. E. J. Brill: 1-958. Herrich-Schaeffer, G. A. W. 1850. Die Wanzenartigen Insekten 9:181-217. C. H. Zeh’schen Buchhandlung, Nurnberg. Hussey, R. F. 1927. On some American Pyrrhocoridae (Hemiptera). Bull. Brooklyn Entomol. Soc. 22(4):227-235. Hussey, R. E and E. Sherman. 1929. General Catalogue of the Hemiptera. Fascicle III Pyr- rhocoridae. Publ. Smith College, Northampton, MA, USA: 1-144. Mayr, G, L. 1865. Diagnosen neuer Hemipteren. Verb. Zool.-Bot. Ges. Wien. 15:429-446. Perty, M. 1830-1834. Delectus animalium articulatorum, quas in itinere per Brasiliam annis 1817-1820, jussu et auspiciis Maximiliana Joseph! Bavariae regis augustissimi peracto, colegerunt Dr. J. B. de Spix et Dr. C. F. Ph. de Martins. Monachii: 1-224. Schmidt, E. 1931. Zur Kenntnis der Familie Pyrrhocoridae Fieber. (Hemiptera-Heteroptera). Stett. Entomol. Ztg. 92(1): 1-51. Stal, C. 1860. Bidrag till Rio Janeiro-traktens Hemipter-Fauna. Kongliga Svenska Vetenskaps- Akademiens Hand, 11(7): 1-84. Stal, C. 1862. Hemiptera Mexicana enumeravit speciesque novas descripsit. Stettin. Entomol. Ztg. 23:287-325. Walker, F. 1873. Catalogue of the specimens of Hemiptera Heteroptera in the collection of the British Museum. Part VI.: 1-210. Received 7 February 1997; accepted 21 June 1997. J. New York Entomol. Soc. 105(1-2): 15-23, 1997 A NEW SUBGENUS AND TWO NEW SPECIES OF THE GENUS MACHILINUS (MEINERTELLIDAE, ARCHAEOGNATHA = MICROCORYPHIA, “APTERYGOTA”, INSECTA) FROM MEXICO Helmut Sturm Department of Biology, University of Hildesheim, D-31141 Hildesheim, Germany Abstract. — Protomachilinus NEW SUBGENUS, M. (P.) mexicanus NEW SPECIES and M. (P.) oaxacensis NEW SPECIES are described. The new subgenus is more plesiomorphic than any of the other three subgenera of Machilinus. Ecological and biogeographical aspects of the genus are discussed. The genus Machilinus has a special position within the Meinertellidae, a family which is more derived than the other family of extant Archaeognatha, the Machilidae. There are three reasons for this. From the morphological point of view the Mach- ilinus species described up until now are highly derived, e.g. by sublateral ocelli which are round to elliptical and hyaline to light red; the absence of a horizontal process on the base of the maxillary palp (a process which is present in all other genera of Meinertellidae); the absence of coxal stylets on all legs; the absence of coxal vesicles on abdominal coxostemite I and the highly reduced chaetotaxy on gonapophyses VIII and IX of the females (Sturm and Bach, 1993). The ecology of this taxon also shows some peculiarities. Contrasting with nearly all other genera most Machilinus species are active during full daylight (The only other species for which daylight activity is described is Bachilis multisetosa of the family Machilidae: Mendes, 1977). Some species were found in very dry localities and one species collected at 4500 m above sea level, near Tucuman, Argentina (Wygodzinsky, 1967). Noteworthy is also the biogeographical aspect. The species of the genus described since then were collected in the western part of North Amer- ica, the southern part of South America, the mediterranean area, the middle of the Sahara and in South Africa. As a result Machilinus is second only to the genus Machiloides (Meinertellidae) in being the most widespread genus of the Archaeog- natha (Wygodzinsky, 1967 Sturm, 1984). The present work reports a new subgenus from Mexico. This finding takes on added importance by the fact that this subgenus shows several plesiomorphic char- acters and is doubtless the most primitive of the four subgenera of Machilinus. Some of its characteristics establish a connection between the genus Machilinus fairly isolated before and the other genera of the Meinertellidae. These characters also make it necessary to redescribe some generotypic features. As a result of the dis- covery of the new subgenus in Mexico questions arise about biogeography and evolution of the genus, discussed after the description. Protomachilinus Sturm, New Subgenus Type species: Machilinus (P.) mexicanus, new species Diagnosis: As subgenus Machilinus s.str. except: Lateral ocelli big, elliptic, with 16 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) black pigment situated on an indentation of the ventral margin of the eyes; 4 distal teeth of mandible without distinct pigment. — Maxillary palps: Horizontal projection on the dorsal basis present; article 2 of male without distinct specializations referring to form and chaetotaxy. — Labial palps: Article 3 distinctly broadened, in males more than in females. — Abdomen: Sternocoxites I with 1 + 1 functioning coxal vesicles. Etymology: The subgenus is named after the accumulation of plesiomorphic char- acters: protos (Greek) = first (in the sense of original). Discussion: The combination of characteristics mentioned above distinguish the sub- genus from the 3 other subgenera of the genus. They are (with exception of the broadened article 3 of labial palps) plesiomorphic and are present either in all other genera of Meinertellidae (horizontal process on the basis of maxillary palps) or the majority of the taxa (lateral ocelli with distinct black pigment, teeth of mandible without pigment, coxal vesicles on stemocoxite I present). Certainly the interrupted pattern of macrochaetae on gonapophyses VIII and IX of female (ovipositor-type IV after Sturm and Bach, 1993) is present but it is more weakly developed than in the other 3 subgenera (see Figs. 25, 26). On the other hand in view of the number of differences with the other subgenera the question arises if this taxon could should actually be a genus. The main reason against this possibility are the many derived common characteristics, shared with the other subgenera of the genus Machilinus (Sturm and Bach, 1992), e.g. sublateral position of lateral ocelli and their ovoid or elliptical form, size and form of eyes, similar distribution of spiniform setae on the margins of thoracic tergites I-III (Sturm and Messner, 1995); absence of coxal stylets on legs, presence of characteristic setae near the basis of abdominal stylets; penis with thin straight setae on characteristic bumps; derived chaetotaxy of female gonapophyses. Machilinus (Protomachilinus) mexicanus Sturm, New Species Types: Holotype male, body length 7.5 mm; MEXICO, Chiapas, 5 mi W San Cris- tobal, pine-oak-forest, 24. VIII. 1966, J. & W. Ivie coll. — Allotype female, 8.5 mm, same data as type. Paratypes: 1 female 9.5 mm; 1 juvenile 4.5 mm, same data as type. All specimens deposited in AMNH (3 adult specimens prepared on slides: Euparal inclusion). Etymology: the species is named after the country of origin, Mexico. Description: Body length ca. 7-10 mm; hypodermal pigment dark brown, extended especially on head, mandibles, maxillary palps, labium, legs and thoracic stemites (see Figs. 1, 4, 6-9). Head (Fig. 1): Eyes large; ratio of width of eyes : width of head = 0.75-0.83; length of eyes : width ca. 0.8; line of contact of the eyes : length of eyes = 0.6-0.7. — Paired ocelli sublateral; contrasting with the other subgenera by black pigment and great size; width reaching nearly half width of eyes. — Antennae (Figs. 2, 3) shorter than body, up to 6.5 mm long, with weak and uniform light brown pigmentation; scapus short, ratio length to width == 1.5-1. 9, distal chains of the flagellum with up to 11 articles, obviously limited by jointlets. — Mandibles distally with 4 teeth of equal size, without distinct pigmentation. — Maxillary palps (Figs. 4, 5): On the dorsal base a horizontal process is present; such a process is absent in all other subgenera of Machilinus but present in all other Meinertellidae; hook on article 2 of male and 1997 MACHILINUS FROM MEXICO 17 chaetotaxy on articles 2 + 3 without distinct peculiarities; number of spines on articles 1! 6! 5 = ca. 10/ 13/ 3; ratio length of articles 7:6:5:4;3 == 1:1.7-2:1.85- 2.05:1-1.1:0.9-1 (for the 2 females), articles 4-7 in the male absent (broken). — Labium (Figs. 6, 7): Article 3 of palps distally distinctly broadened, in the male more than in the female. Thorax: Number of lateral marginal spiniform setae on tergites I/II/III = 12-15/35- 47/10-16 (cf. Sturm and Messner, 1995). — Legs (Figs. 8, 9): Coxal stylets absent; some dark spiniform setae on tarsal articles 1 + 2 of legs I. Abdomen: Coxites I-VII each with 1 + 1 functioning coxal vesicles, II-IX with stylets; all stemites small, terminal spines of stylets long (Figs. 12, 13), ratio length of terminal spine: length of stylet for segments II = 0.4-0. 5:1 for V == 0.8-l,0:l; for IX = 0.6-0.65:1 (determined only in male); distance between inner margins of basis of abdominal stylets only slightly different, ratio of distance segment IV:II = 1.2-1.35 (cf. description of the other subgenera in Sturm and Bach, 1992); coxites II + III with small fields of setae lateral to the base of stylets, setae up to ca. 0.15 mm long, straight and of brown colour (Fig. 11); stylets IX ventrad and mediad with brown spiniform setae, up to 0.45 mm long (Fig. 13). — Penis (Fig. 14): distinctly shorter than 0.5 X length of coxite IX; aperture pointed oval, surrounded by 1-3 rows of straight setae (up to 0.03 mm long) rising on small characteristic bumps. Ovipositor extended far beyond terminal spines of stylets IX; gonapophyses with more than 60 articles, only on the distal third with macrochaetae; terminal spine of middle size, reaching the length of 2-3 terminal articles; 3-5 setae per article on the four most distal articles, proximadly reduced to 1—3; lateral macrochaetae in this latter area not present on each article (interrupted pattern of distribution after Sturm and Bach 1993). — Cerci with long straight terminal spines (Fig. 18). Diagnosis: The species can be determined by the characteristics of the subgenus and the differences which are given in the diagnosis of Machilinus (P.) oaxacensis. Machilinus (Protomachilinus) oaxacensis Sturm, New Species Types: Holotype male, body length 6.5 mm; MEXICO, Oaxaca county, near Tla- colula, 96.27 W, 16.57 N, under cliff, 30.IV. 1963, Gertsch & Ivie coll.— Allotype: female, body length 7 mm, same data as holotype; paratypes 1 male, 4 females, same data as holotype. — Paratypes: 2 females, 6.5 and 8 mm; MEXICO, Oaxaca, 3 mi SE Tlacolula 96.25 W, 16.56 N, 30 VIII. 1966, J.& W. Ivie coll. — All types deposited on AMNH, 6 adult specimens prepared on slides (Euparal inclusion). Etymology: The species name refers to the province Oaxaca (Mexico) where the species was collected. Description: Body length (inclusive of head) ca. 6-8 mm; hypodermal pigment dark brown, more pronounced than in the first species. Head: Hypodermal pigment on frons V-shaped, with small median stripe (Fig. 19). — Ratio of width of eyes: width of head == 0.75-0.85; length of eyes: width = 0.75- 0.9; length line of contact of eyes: length of eyes = 0.55-0.65; width of ocelli : width of eyes = 0.45-0.55 (n = 6 for all values); ocelli black. Antennae distinctly shorter than the body, maximum length measured = 3.5 mm; maximum number of articles per chain within the distal half = 9; patches of hypodermal pigment on scapus and pedicellus only; flagellum uniformly light brown; scapus short, ratio length : width 18 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) = 1. 5-1.9 — Maxillary palps especially on article 3 stronger pigmented than in the first species; setae on the medial side of article 2 forming an indistinct whorl (Fig. 20); ratio length of articles 7:6:5:4:3 = 1:1.4-1.6:1.7-2.3:1.1-1.3:1.1-L5 (n = 12); number of spines on articles 7/6/5 = 5-7/9-11/1-2. — Broadening on article 3 of labial palp of female weak, of male very distinctive. Thorax: Legs more intensive and more extended pigmented than in the first species; spiniform setae on all legs present (Fig. 22); maximum numbers on femur/tibia/tarsus 1/tarsus 2 for leg I = — Z3/3/5; for II = 2/5/4Z4; for III = 3/5/4Z4. Maximum number of lateral marginal setae on tergites I/II/III = 19 (partially in double rows)/>50/ 16. — All stemites with big patches of hypodermal pigment. Abdomen: Coxal vesicles on coxites I-VII; stylets on II-IX; coxites I with a small band of setae on the anterior border (Fig. 23); fields of setae near the base of stylets on coxites II- VIII present; terminal spines on stylets long; ratio of length of spine : length of stylet on segment II ca. 0.5, on V ca. 0. 5-0.6, on IX ca. 0.35-0.45. Penis: Aperture great, pointed triangular; structure and taxis of setae similar to those of the first species. Ovipositor thin, extended well past the ends of terminal spines IX; chaetotaxy similar to the first species. Filum terminate and cerci with many dark brown spiniform setae; cerci up to 3.3 mm long with one terminal spine of medium size. Diagnosis: Machilinus (P.) oaxacensis is distinguished from the other species of the subgenus e.g. by the shorter body length, the different pigment pattern on the frons and maxillary palps, different ratios of length of articles on maxillary palps, the strong broadening of article 3 on the labial palps of the male, presence of dark spiniform setae on the tibia and tarsus of all legs, and the presence of fields of setae near the bases of stylets II- VIII. Discussion: The species presents all characteristics of the subgenus. Specimens which were collected in 1966 at a locality near the type locality included a female (9 mm body length) with a distinctly different ratio of distances between stylets on segments II and IV {= 1,14) and some other peculiarities. It can not put to this species but indicates that there are probably more species of the subgenus which contrasts with the subgenus Nearctolinus with only one species and a much larger distribution. Figs. 1-18. Machilinus {Protomachilinus) mexicanus, new species. 1. Head (male), frontal view. 2. + 3. Antenna, female; 2. Basal part with scapus + pedicellus; 3. Distal part, ca. 4 mm from base. 4. Maxillary palp (female), lateral view. 5. Articles 1-3 of male maxillary palp, median side, article 2 with hook. 6. Left half of labium with labial palp (male), ventral view. 7. Labial palp (female), articles 2 + 3.8. Leg I (male). 9. Leg III (male). 10. Half of coxosternite I (male), ventral view. 11. Part of coxite III (male) with base of stylet and lateral setal area, ventral view. 12. Half of coxosternite V (male) with muscles (dotted) and coxal vesicle, ventral view. 13. Coxite IX (male) with penis, ventral view. 14. Penis of Fig. 13 by higher magnifi- cation. 15. + 16. Distal parts of gonapophyses VIII (female), ventral view. 17. Distal part of gonapophysis IX (female), ventral view. 18. Distal end of cercus (female) with big terminal spine. 1997 MACHILINUS FROM MEXICO 19 0.05 mm 20 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Figs. 19-26. Machilinus {Protomachilinus) oaxacensis, new species; hypodermal pigment dotted. 19. Head (male), frontal view. 20. Maxilliary palp (male), articles 1-5, lateral view, inner side. 21. Left half of labium (male) with labial palp, ventral view. 22. Distal part of leg III (male) with spiniform setae. 23. Half of sternocoxite I with coxal vesicle and fields of setae, ventral view. 24. Penis, ventral view. 25. Distal part of gonapophysis IX (female), ventral view. 26. Distal part of gonapophysis IX of Machilinus (Nearctolinus) aurantiacus: Interrupted pattern of macrochaetae more significant than in 25.; ex Sturm and Bach 1992. 27. Machilinus {Neo- tropolinus) chilensis Sturm and Bach, 1992: Maxillary palp (male), articles 2 + 3: high spe- cialized chaetotaxy and form of article 2; ex Sturm and Bach 1992. ECOLOGICAL AND BIOGEOGRAPHICAL ASPECTS OF THE GENUS MACHILINUS The localities where specimens of the genus were found show very different eco- logical conditions: rocks or stones of very different composition, sand dunes, bark of fallen trees, litter, heath and grassland. The range of altitude reaches from marine beaches (Mendes, 1977) up to 4500 m above sealevel (Argentina: Wygodzinsky, 1967). Peculiar for Archaeognatha is the activity in full light. It was described by different authors (e.g. Wygodzinsky, 1952, Bitsch, 1968, Smith, 1970, Mendes, 1977) and is probably valid for most of the Machilinus species. For nearly all other 1997 MACH I LINUS FROM MEXICO 21 genera activity during night or dusk is regular. Only for Bachilis multisetosa (Mach- ilidae) Mendes (1977) mentioned a less pronounced daylight activity. Perhaps the light or reddish colour described for the three other subgenera of Machilinus and for Bachilis m. is connected with this form of activity. The black colour of the paired ocelli in the subgenus Protomachilinus corresponds to that of nearly all other Ar- chaeognatha and it remains unclear if it indicates activity in darkness or twilight. The hygrophily is apparently developed differently within the genus. Besides species or populations which are living in habitats with high relative humidity (e.g M. chi- lensis in the forests of Chile; (Sturm and Bach, 1992) or M. gredosi in the Serra da Estrela, Portugal, an area with 1900 mm precipitation per year: Mendes 1977) there are some species living in dry areas (e.g., M. aurantiacus on the sand dunes near Los Angeles [collected by R. Mattoni, Beverly Hills: VII. + VIII. 1993 in pitfall traps] and on dry localities in Utah and New Mexico (Mendes, 1996) or active for long time in full sunlight. In the latter cases a regular production of dew during the night could stabilize the water balance. But it is an enigma why M. aurantiacus often living in dry habitats would have strongly reduced coxal vesicles. For other bristle- tails the uptake of water by means of the coxal vesicles is regular (Bitsch, 1974). It may be that the genus Machilinus in connection with the daylight activity has achieved other means of water balance. The genus Machilinus does not enter tropical lowlands. The collecting localities in Mexico are lying at altitudes of more than 1000 m, those at Tibesti Mountains and in Yemen at more than 2000 m. The geographic distribution of the genus Machilinus is characterized by a multi- tude of isolated and widely distributed areas. Three of these areas are situated in America, each harboring one subgenus. The number of species decreases from South to North with South America having 6, Mexico 2, North America 1 (Fig. 28). The forth subgenus is restricted to the Western part of the Old World and distributed in at least 4 clearly separated areas (at the basis of the data known at the present time). Old data referring to the occurrence on the Cape Verde Islands (Silvestri, 1908) could not be confirmed. The area with the highest number of species (18) is the Mediterranean. The remaining areas are very small, each harboring 1 species. The occurences in Tibesti, Yemen and South Africa could be relicts, that in Madeira perhaps a result of dispersal. Within the four subgenera of Machilinus, Protomachilinus undobtedly shows the most plesiomorphic characters, which makes the genus morphologically more similar to the other genera of Meinertellidae. An undescribed specimen from the Lebanese amber (collection of the American University at Beirut, number 194/35) gives evidence for the existence of typical representatives of the family Meinertellidae some 110-120 millions of years ago. The splitting within this family must have been happened distinctly earlier, perhaps at the beginning of the Mesozoic. An explanation for the present distribution on the basis of the known historical, biogeographic and morphological data remains diffi- cult, but a comparison with the distribution of another taxon of the Meinertellidae, the genus Machiloides could be helpful. Machiloides is not only the most plesio- morphic genus in the Meinertellidae (e.g. coxal stylets on legs II and III, lateral ocelli sole-shaped, chaetotaxy on genital appendages not specialized) but it shows also the most extensive distribution within the extant Archaeognatha (Table 1). 22 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) hatched; numbers = number of species per area; ? = occurrence uncertain. The fact that no other genus of Archaeognatha shows such an extensive distribution as these two genera suggests the long period of time necessary to reach such a wide distribution and morphological differentiation. Probably the origin of both genera and of all Archaeognatha was in the northern hemisphere, where presently occur the most plesiomorphic species of Machilinus and Machiloides (Machilinus [Protomachilinus] in Mexico and Machiloides tenuicomis in Spain). If Laurasian in origin, the dispersal of Machiloides began earlier than that of the more derived genus Machilinus, which ap- parently never reached the Austral-malesian area. A similar pattern of distribution is in the genus Araucaria (Gymnospermae) during the Mesozoic, especially during the Cre- Table 1. Distribution and number of species for each region for the genera Machilinus and Machiloides (Meinertellidae). Machilinus Machiloides North America 1 2 Central America 2 — South of South America 6 10 South and East Africa 1 25 Mediterranean Region 19 1 Australia/Tasmania — >2 1997 MACHILINUS FROM MEXICO 23 taceous period, and included North America, Southern South America, Europe, South Africa, India and the Australian region (Ehrendorfer, 1978). ACKNOWLEDGMENTS I am indebted to Dr. Randall T. Schuh of the American Museum of Natural History, New York for the loan of this interesting material, to Dr. Grimaldi of the same Museum for the revision of the English text. LITERATURE CITED Bitsch, J 1968. Les especes circum-mediterraneenes du genre Machilinus (Insecta Thysanura). Bull. Soc. Sci. Nat. et Phys. du Maroc 47:165-192. Bitsch, J. 1974. Eonction et ultrastructure des vesicules exsertiles de 1’ abdomen des Machilides. Pedobiologia 14:142-143. Ehrendorfer. E 1978. Geobotanik. In: Denfer, v.D. et al. (eds.): Lehrbuch der Botanik fiir Hochschulen. G. Eischer V., Stuttgart: 948. Mendes, L. F. 1977. Notations chorologiques sur les Microcoryphia du Portugal. Rev. Ecol. Biol. Sol 14(l):63-70. Mendes, L. F. 1996. Some new data on the Microcoryphia and Zygentoma (Insecta) from the United States. Garcia de Orta, Sen Zool. Lisboa 21(1):1 17-126. Silvestri, F. 1908. Tisanuri raccolti da L. Fea alle isole del Capo Verde . . . Ann.Mus. Stor. nat. Genova (3)4:137-187. Smith, E. L. 1970. Biology and structure of some California bristletails and silverfish. Pan- Pacific Entomologist 46:212-225. Sturm, H. 1984. Zur Systematik, Biogeographie und Evolution der siidamerikanischen Meiner- tellidae (Machiloidea, Archaeognatha, Insecta). Z.Zool. Syst. Evolut.-fschg. 22:27-44. Sturm, H. and C. Bach de Roca. 1992. New American Meinertellidae (Archaeognatha, Mach- iloidea). Pan-Pacific Entomoglo. 68(3):174-191. Sturm, H. and C. Bach de Roca. 1993. On the systematics of the Archaeognatha. Entomol. Gener. 18(1/2): 55-90. Sturm, H. and B. Messner. 1995. Chaetotaxy of tergites in the Archaeognatha (Insecta) and its taxonomic significance. Zool. Anz. 234:85-100. Wygodzinsky, P. 1952. Apuntes sobre “Thysanura” americanas (Apterygota, Insecta). Acta Zool. Lilloana 11:435-458. Wygodzinsky, P. 1967. On the geographical distribution of the South American Microcoryphia and Thysanura (Insecta). In: Delamare Deboutteville, Cl. and E. Rapoport: Biologic de I’Amerique Australe vol. 111:505-524, Paris. Received 24 February 1997; accepted 5 August 1997. J. New York Entomol. Soc. 105( l-2):24-39, 1997 A REVIEW OF THE GENUS LIMNOMETRA MAYR IN NEW GUINEA, WITH THE DESCRIPTION OF A VERY LARGE NEW SPECIES (HETEROPTERA: GERRIDAE) Dan a. Polhemus' and John T. Polhemus^ 'Dept, of Entomology, MRC 105, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 -Colorado Entomological Museum, 3115 South York Street, Englewood, CO 80110 Abstract. — The species of the water strider genus Limnometra occuring on New Guinea and proximate islands are reviewed, and a very large new species, L. grallator, is described from the southern flanks of the central highlands. On the basis of male endosomal sclerotization, this new species appears to be most closely related to L. kallisto (Kirkaldy), a much smaller species also endemic to greater New Guinea. A key to all New Guinea Limnometra species is provided, accompanied by illustrations of the male endosomal sclerites and range maps. The genus Limnometra consists of a group of large, long legged water striders that are widely distributed across tropical Asia and eastward into the southwestern Pacific and northern Australia. The last complete generic revision of the genus was that of Hungerford and Matsuda (1958), following which no additional species were described for over 40 years. Recently, however, Nieser and Chen reviewed the Indo- Australian fauna (1992), describing 9 new species and producing a partial key. Sub- sequently, Andersen and Weir (1997) revised the genus for Australia, describing one new species, L. cilioides, keying the Australian taxa, and correcting certain errors contained in Nieser and Chen, including synonymy of their L. poliakanthina from Queensland. While Andersen and Weir’s work was in press, the first author (DAP) collected examples of still another very large new species from New Guinea, de- scribed herein as L. grallator, during the course of biological surveys in the Kikori Basin of southern Papua New Guinea and the Timika area of southern Irian Jay a. This latter species is the largest known from New Guinea, and it seems rather ironic that it was the last to be collected. This spate of recent taxonomic work and collecting has necessitated the current review of Limnometra in New Guinea. The present paper includes a key to the 5 species known to occur on the island, a description of L. grallator, and distribution maps and records for all species treated. The geographical coverage includes New Guinea proper, and certain proximal islands (Salawati, Biak, Yapen, Woodlark, Normanby), but excludes the Bismarcks and Solomons, where certain unresolved taxonomic problems remain. Distributional information for taxa occurring in the Bismarcks is thus included only for those species also occurring on New Guinea proper. Of the 5 species now known to occur on New Guinea, three, L. kallisto (Kirkaldy), L. ciliodes Andersen and Weir, and L. lipovksii Hungerford and Matsuda, are widespread within the island. The remaining two species are more circumscribed in their distributions; L. ciliata Mayr is found only in the northern half of the island and in the Papuan Peninsula, areas which represent geological terranes accreted to New Guinea from the Miocene onward, while L. grallator 1997 LIMNOMETRA OF NEW GUINEA 25 n. sp. is found only south of the central mountains on portions of the island that are part of the original Australian craton. Since Limnometra are large and obvious, they are captured by general collectors more often than other aquatic Heteroptera. As a result, a large historical specimen base exists, particularly at the Bishop Museum in Honolulu. These previous collec- tions have been further augmented over the last five years through surveys by the authors in poorly known sections of Irian Jaya and southern Papua New Guinea. The maps presented herein thus provide relatively detailed depictions of species distributions within the island. All measurements are given in millimeters. CL numbers following certain local- ities refer to a coding system used by the authors to cross-reference specimens and collection data. KEY TO SPECIES OF LIMNOMETRA OCCURRING ON NEW GUINEA 1. Connexival spines not reaching beyond apex of genital segments; eighth abdominal segment (first genital segment) of male with a backward pointing tooth-like projection on either side ventrally L. lipovskyi Hungerford and Matsuda - Connexival spines reaching beyond apex of genital segments (Fig. 3); eighth abdom- inal segment (first genital segment) of male without a backward pointing tooth-like projection on either side (may be produced laterally, but not in a tooth-like projection) 2 2. Middle and hind femora dark on basal half, distinctly pale yellowish distally; only macropterous form present kallisto (Kirkaldy) - Legs unicolorous; both macropterous and reduced winged forms may be present 3 3. All legs uniformly pale and straw colored; connexival spines very long, curved from base (Fig. 3); mid femora without long cilia grallator D. and J. Polhemus - All legs dark colored; connexival spines shorter, straight; mid femora with long cilia 4 4. Eighth abdominal segment (first genital segment) of male produced laterally on either side ventrally; macropterous forms with medial dark stripe on pronotum bordered to either side by pale stripes that increase in width posteriorly, these stripes often becom- ing greatly expanded to form broad pale patches that cover most of the pronotal disk; paired dark stripes on mesopleura always well defined, usually with an area of pale whitish coloration between them ciliata Mayr - Eighth abdominal segment (first genital segment) of male not, or only very weakly, produced laterally on either side ventrally; macropterous forms with medial dark stripe on pronotum bordered to either side by narrow pale stripes of similar width to the medial stripe, these pale stripes retaining an even width throughout their length, not becoming expanded posteriorly; paired dark stripes on mesopleura often obscure or absent, if present then usually lacking a pale whitish area between them cilioides Andersen and Weir Limnometra grallator, n. sp. Figs. 1-6, 11 Diagnosis: Recognized by its large size (body length in brachypterous males ex- ceeding 19.0 mm), yellowish-brown ground color with sharply contrasting black stripes (Fig. 1), straw colored legs without dark markings, and asymmetrical, V- shaped ventral endosomal sclerite in the male (Fig. 6). 26 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Figs. 1-3. Limnornetra grallator n. sp. 1. Male, lateral habitus; legs omitted. 2. Male pro- notum, showing pattern. 3. Male terminal abdomen, dorsal view, showing length and shape of connexival spines. Description: Size. Macropterous male, length 17.0-17.2, body width 4.9-5. 1, head width 2. 7-2. 8; brachypterous male, length 19.0-21.5, body width 6. 0-6. 5, head width 2. 7-2. 8; macropterous female, length 16.5—17.0, body width 4. 0-4. 5, head width 2.4— 2.5; brachypterous female unknown. Color. Head and pronotum yellowish brown in ground color with very distinct black or dark brownish markings (Figs. 1, 2). Head between eyes with median pair of indistinct longitudinal stripes, these stripes divergent anteriorly, meeting posteri- orly; a well defined dark stripe also present along inside margin of each eye; antennae brown, unicolorous, without white markings. Pronotum with relatively narrow, black, longitudinal median stripe bordered by broad pale bands, this median stripe of rel- atively uniform width throughout. Lateral and posterior portions of pronotum bearing black stripe parallel to margin, this stripe wider posteriorly; entire margin of pronotal lobe outside of this black stripe narrowly yellow (Fig. 2). Thoracic pleura yellowish; propleuron with broad black band behind eye; mesopleuron with two longitudinal black stripes, the upper one continued as an irregular dark stripe on the metapleuron. 1997 LIMNOMETRA OF NEW GUINEA 27 Proacetabulum with an irregular dark spot posteriorly; meso- and metacetabulae each with a dark stripe dividing two pale areas (Fig. 1). Fore wings medium brown to orange brown, with sharply contrasting dark brownish or blackish veins, but without pale subcostal markings. Fore femur and tibia uniformly yellowish brown, infuscated ventrally, fore tarsi dark brown; middle and hind legs uniformly pale yellowish brown, without pale annulations or dark apical markings. Abdomen (when visible in brachypterous form) black dorsally, with connexival margins narrowly yellow; con- nexival spines dark. Ventral surfaces of head, thorax and abdomen pale yellowish. Brachypterous male elongate, length 3.4X greatest body width (17.2:5.1). Head between eyes (anterior interocular) 1.7X longer than wide (2.0: 1.2); lengths of an- tennal segments I-IV: 5.3, 3.7, 4.5, 5.0; third and fourth rostral segments together about 1.3X maximum head width across eyes (4. 1:3.1). Thorax with pronotum long (7.9), anterior part 0.8 X wide as head (2.7:3. 1), pronotal lobe with slightly divergent sides, widest at humeri (3.2), posterior margin angulate; meso- and metastemum with pale recumbent pubescence. Leg segments with lengths as follows (femur: tibia: tarsal 1: tarsal 2): fore leg, 8.5, 7.0, 1.2, 1.9; middle leg, 37.5, 38.0, 6.0, 1.0; hind leg, 39.0, 38.0, 2.3, 0.8; fore femora straight and relatively slender, maximum width distinctly less than width of middle femora (0.4:0.6); fore tibia straight; middle femur densely clothed with minute, slender, black, spine-like setae ventrally, and bearing 6-10 short, stout, black, tooth-like spinules distally, but lacking any fringe of long hairs; middle tibia similarly setiferated, bearing a longitudinal row of tiny, slender black spines along entire length; setiferation of hind femur and tibia similar to that of middle leg, but hind femur lacking tooth-like spinules distally. Wings narrow, only partially developed, extending half the length of abdomen Abdomen long and nearly parallel-sided, widening slightly posteriorly, maximum width about 0.27 X total length along dorsal midline (2.7:10.0); lengths of abdominal tergites IV-VI (only segments visible beyond wings) as follows: 1.0, 1.0, 1.2. Connexival spines very long, parallel to slightly divergent and basically horizontal, curving from bases, well over 2X as long as the visible genital segments when viewed from above (Fig. 3) ; abdominal venter distinctly carinate, stemite VII shorter than stemite VI, with hind margin simply concave. Genital segments relatively small; venter of segment VIII distinctly shorter than that of segment VII; pygophore parallel sided, rounded distally; parameres small, rounded distally, sparsely set with extremely short setae; proctiger conical distally; vesical armature relatively simple; dorsal plate not well defined; dorsal scelerite robust, symmetrical, with distinctly widened and curved short distal part, basal part divided (Fig. 5); lateral sclerites extremely large and long (Fig. 4) ; accessory sclerites not evident; ventral sclerite V-shaped, asymmetrical (Fig. 6). Macropterous female considerably smaller than brachypterous male, but otherwise similar in general structure and coloration, with following exceptions: forewings long, reaching to tips of connexival spines, coloration brown with darker veins, lacking pale subcostal markings; gonocoxae relatively small, exposed ventrally; proc- tiger small, elongate, semi-acuminate. Macropterous male: Similar to brachypterous male in general structure and col- oration, but smaller in overall body dimensions (see above). Wings medium brown, with slightly darker brown venation, lacking pale subcostal markings, extending to tips of abdominal connexival spines. Brachypterous female: Unknown. 28 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Figs. 4-6. Male endosomal structures of Limnornetra grallator n. sp. 4. Lateral view. 5. Dorsal view. 6. Ventral view (note asymmetrical, V-shaped ventral sclerite). Figs. 7-10. Male endosomal structures of Limnornetra species occurring on New Guinea (all shown in lateral view). 7. Limnornetra ciliata Mayr. 8. Limnornetra ciliodes Andersen and Weir. 9. Limnornetra kallisto (Kirkaldy). 10. Limnornetra lipovskii Hungerford and Matsuda. 1997 LIMNOMETRA OF NEW GUINEA 29 Table 1. Comparison of external structural characters between Limnometra grallator n. sp. and Limnometra ciliodes Andersen and Weir: Character ciliodes grallator Leg color Dark brown Yellow Setae on middle femur Long Short Length of legs Long Extremely long Ratio of antennal segments II/IV 25/18 (male) 23/38 (male) Color of figures on head vertex Black Brown Macropterous pronotal margins Broadly dark Narrowly dark Mesopleural coloration Light, or with two With two strong black modest stripes stripes Connexival spines Long, straight, and slightly Extremely long, curved divergent distally from base Etymology: The name grallator (Latin, masculine), means “one who walks on stilts”, and refers to the extremely long legs of the brachypterous males in this species. Discussion: Among New Guinea Limnometra species, L. grallator n.sp. is superfi- cially most similar to L. ciliodes Andersen and Weir, but may be separated by the external characters given in the comparison table above. The structure of the endomosal sclerites is also very different between the two species (compare Figs. 4 and 8). In particular, L. grallator has an asymmetrical, V- shaped ventral sclerite (Fig. 6) quite unlike the symmetrical, U-shaped ventral sce- lerite of L. ciliodes, but similar to the state seen in L. kallisto. The endosoma of L. ciliodes also bears several accesory lateral sclerites similar to those seen in L. ciliata (compare Figs. 7 and 8); such accessory sclerites are lacking in L. grallator and L. kallisto (Figs. 4, 9). Based on current records, L. grallator is confined to the premontane foreland of the central mountain ranges in southern New Guinea, from the Kikori River basin westward to at least the Timika area (Fig. 11). Individuals prefer small, shaded forest streams, where they skate on sheltered pools, often occurring sympatrically with L. kallisto. In contrast to most Limnometra species, in which macropterous morphs predominate, the macropterous forms of L. grallator appear to be very uncommon, with only a few examples currently known. Distribution: Southern New Guinea (Fig. 11). Material examined (all specimens brachypterous unless otherwise noted): HOLOTYPE, brachypterous male, PAPUA NEW GUINEA, Southern Highlands Prov., branch of upper Kara Creek (trib. to Digimu River), 5.5 km. S. of Moro oil camp, on road to Ridge camp, 900 m., water temp. 20°C., 12 March 1995, D. A. Polhemus (BPBM). PARATYPES: PAPUA NEW GUINEA, Gulf Prov.: 2 males, 1 female (macrop- terous), Omo River at Omo, 40 m., water temp. 24°C., 28 February 1995, D. A. Polhemus (BPBM); 1 male (macropterous), Lubu River above Omo, 50 m., water temp. 19°C., 1 March 1995, D. A. Polhemus (BPBM); 1 male, 3 females, Wamivei Creek, near Veiru Creek, Kikori River delta, SW of Kikori, 40 m., water temp. 25°C., 5 March 1995, D. A. Polhemus (BPBM). Southern Highlands Prov.: 15 males, 8 females, same data as holotype (BPBM, USNM, JTPC); 2 males, 1 female, branch 30 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Fig. 11. Distribution of Limnometra grallator n. sp. in New Guinea. Stippling indicates areas above 2000 m. of upper Kara Creek (trib. to Digimu River), 3.5 km. S. of Moro oil camp, on road to Ridge camp, 900 m., water temp. 20°C., 13 March 1995, D. A. Polhemus (BPBM); 1 male, 1 female, small rocky creek in disturbed forest, approx. 1.0 km. N. of Tubage, NE of Moro, 1000 m., water temp. 21°C., 14 March 1995, 14:30-15:00 hrs.; 22 March 1995, D. A. Polhemus (BPBM); 1 male, small rocky creek approx. 1.25 km. N. of Tubage, NE of Moro, 1000 m., water temp. 20°C., 14 March 1995, 15:00-15:30 hrs.; 22 March 1995, D. A. Polhemus (BPBM); 1 male, rocky stream at Ludesa Mission, N. of Mt. Bosavi, 700 m., water temp, of stream 23°C., 20 March 1995, D. A. Polhemus (BPBM). Western Prov.: 1 male, 2 females, Olsobip, Fly River, 400-600 m., August 1969, J. and M. Sedlacek (BPBM). INDONESIA, Irian Jay a Prov.: 2 males, 1 female. Tributary to Iwaka River, ap- prox. 3 km. W. of Kuala Kencana, N. of Timika, 4°24.08 S, 136°50.10 E, 300 ft., water temp. 25°C., pH 6.95, 17 January 1997, 09:30-12:00 hrs., CL 7042, D. A. Polhemus (NMNH, LIPI); 6 males, 7 females, same locality as preceeding except 26 March 1997, D. A. Polhemus, J. T. Polhemus, and T. Tabuni (JTPC, LIPI); 6 males, trib. to upper Minajerwi River, approx. 1 km. E of Mile 50 tank farm on Tembagapura Road, 4°16.99 S, 137°01.56 E, 1650 ft., water temp. 23°C., 24 January 1997, 12:00-14:00 hrs., CL 7061, D. A. Polhemus (NMNH, LIPI). Limnometra ciliata Mayr Figs. 7, 12 Limnometra ciliata Mayr 1865, Verb, zool.-bot. Ver. Wien, 15: 444 (Type, male, Java, NHMW) Discussion: This is the most widespread of all Limnometra species, occurring in lowland habitats from Indochina through the Malay Archipelago to New Guinea, the Bismarcks, the Solomons, Fiji and Guam. 1997 LIMNOMETRA OF NEW GUINEA 31 Fig. 12. Distribution of Limnornetra ciliata Mayr in New Guinea. Stippling indicates areas above 2000 m. Distribution: New Guinea (Fig. 12), Solomons (Guadalcanal), New Britain, Guam, Fiji, Halmahera, Ambon, Bali, Sumatra, Borneo, Celebes, Mindanao, Luzon, Marin- duque, Thailand, Malaysia. New Guinea records: PAPUA NEW GUINEA: Central Prov.: 1 female. Brown River, 5 m., stream pools, 23 October 1960, J. L. Gressitt (BPBM). East Sepik Prov.: 1 male, Brandi School, nr. Wewak, CL 1795, 10 September 1983, D. A. and J. T. Polhemus (JTPC); 1 male, Mandi Creek, E. of Wewak, CL 1797, 10 September 1983, D. A. and J. T. Polhemus (JTPC); 1 male, Yemogu Creek, 2 km. W of Tring, CL 1805, 12 Septem- ber 1983, D. A. and J. T. Polhemus (JTPC); 2 males, 2 females, waterfall nr. Pasam, CL 1798, 10 September 1983, D. A. and J. T. Polhemus (JTPC); 3 females, Ra- mumba, NW of Wewak, CL 1801, 11 September 1983, D. A. and J. T. Polhemus (JTPC). Madang Prov.: 1 male, 1 female, Friedrichwilhelmshafen, 1901, Biro (JTPC). Milne Bay Prov.: 1 male, S. of Alotau, 29 September 1989, R. N. B. Prior (OXUM). East New Britain Prov.: 2 males, 4 females, Kerawat, Gazelle Peninsula, 60 m., 29-31 August 1955, J. L. Gressit (BPBM). West New Britain Prov.: 2 males. West New Britain, Tamari, rainwater tank, 14 January 1989, R. N. B. Prior (OXUM); 2 males, 2 females. West New Britain, Buluma, nr. Dami, rain water pit, 17 January 1989, R. N. B. Prior (OXUM); 1 male. West New Britain, Blalla Apulpu village, freshwater creek, 27 January 1989, R. N. B. Prior (OXUM). INDONESIA: Irian Jaya Prov.: 1 male, Manokwari, T. Barbour (JTPC); 7 males, 6 females, Yapen Is., Kabupaten Waropen, small rocky stream at Kainsami, 22 Oc- tober 1991, CL 2661, R. Ubaidillah (JTPC, BPBM). Limnornetra cilioides Andersen and Weir Figs. 8, 13 Limnornetra cilioides Andersen and Weir 1997, Invert. Taxon., 11 (2): 242 (Type, macropterous male, N. Queensland, Australia, ANIC) 32 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Eig. 13. Distribution of Limnometra ciliodes Andersen and Weir in New Guinea. Stippling indicates areas above 2000 m. Discussion: All previous records of Limnometra cursitans from New Guinea appar- ently represent this species instead. It is a species of the lowlands, with no records from above 50 m. elevation (Fig. 13). Further discussion of this species’ structural characters and relationships, along with a detailed map of its distribution within Australia, is provided in Andersen and Weir (1997). Distribution: Australia, New Guinea (Fig. 13). New Guinea records: PAPUA NEW GUINEA: Gulf Prov.: 6 males, 15 females, roadside ponds in low- land forest, 2 km. S. of Kopi oil camp, N. of Kikori, 20 m., water temp. 24°C., 27 February 1995, D. A. Polhemus (BPBM). Western Prov.: 3 males, 4 females. Fly River above Kiunga, CL 1774, 3 September 1983, D. A. and J. T. Polhemus (JTPC). INDONESIA: Irian Jaya Prov.: 6 macropterous males, Tanamerah, Station 31, creek in rainforest, 13 June 1957, D. F. M. (ANIC); 2 macropterous males, 5 ma- cropterous females, 3 apterous males, 8 apterous females, Merauke, 27 March 1939, R. G. Wind (SEMK); 1 female, Bernhard Camp, 50 m., July 1938, J. Olthof, Neth. Ind.-American New Guinea Exp. (RMNH); 1 female, Teluk Bintuni, Babo, 13 Au- gust 1941, E. Linquist (NC); 1 male, 1 female. River Tor (mouth), 4 km. E. of Hoi Maffin, 4 July 1959, T. C. Maa (BPBM); 5 males, 3 females, swamp forest pond S. of Walio oil field, nr. Kasim, 7 m. (20 ft.), CL 2620, water temp. 30°C., 29 Sept. 1991, J. T. and D. A. Polhemus (JTPC, BPBM). Limnometra cursitans (Fabricius) Gerris cursitans Fabricius 1794, Entomol. Syst. 4: 192 (Type, macropterous male. Nova Hollandia [= Australia], ZMUC) Limnometra cursitans Banks 1909, Phila. J. Sci. 4: 583. Limnometra poliakanthina Nieser and Chen 1992, Tijdschr. Entomol. 135: 23 (Type, 1997 LIMNOMETRA OF NEW GUINEA 33 Eig. 14. Distribution of Limnometra kallisto (Kirkaldy) in New Guinea. Stippling indicates areas above 2000 m. macropterous male, N. Queensland, Australia, OXUM). Syn. by Andersen and Weir 1997, Invert. Taxon. 11: 236. Discussion: Limnometra cursitans has been recorded from New Guinea in the lit- erature, but these records were based on misidentifications, since the true identity of this species was misinterpreted by Hungerford and Matsuda (1958) in their revision of Limnometra, an error subsequently followed by Nieser and Chen (1992). As now reinterpreted by Andersen and Weir (1997), L. cursitans is restricted to Australia, with all previous records of this species in New Guinea being referable to L. ciliodes Andersen and Weir. Distribution: Australia. New Guinea records: none known. Limnometra kallisto (Kirkaldy) Figs. 9, 14 Gerris kallisto Kirkaldy 1899, Ann. Soc. Entomol. Belg. 43:506 (Type, female, My- sol [= Misool Is.], supposed to be in ISNB, apparently lost). Limnometra kallisto Lundblad 1933, Arch. Hydrobiol., Suppl. 12, Tropische Bin- nengewasser 4: 371. Discussion: This is the most common and widespread species of Limnometra in New Guinea, and the only species, other than the highly precinctive L. grallator, to be found regularly at elevations above 100 meters (Fig. 14). It occupies a wide range of habitats, and individuals from different localities can vary greatly in size. In contrast to other New Guinea Limnometra species, which prefer slow waters, L. kallisto is often found in sheltered eddies along the margins of swift upland streams. Adults will fly readily if disturbed or pursued, alighting amid streamside vegetation. This well developed capacity for flight likely accounts for the species’ broad distri- 34 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) bution within New Guinea, and for its tendency to exploit temporary habitats such as rain pools in forest roads. Distribution: Misool (type locality). Am, Morotai, New Guinea (Fig. 14), New Ireland, Solomons (Bougainville, Guadalcanal, Kolombangara, Malaita, San Cristo- val, Vella Lavella, Choiseul). New Guinea records: PAPUA NEW GUINEA: Central Prov.: 1 male, 2 females, 3 km. S. of Vanapa, Brown River road, 200 m., 17-26 May 1965, around forest pool, W. A. Steffan (BPBM); 1 male, Mt. Suckling, 14 June 1972, W. H. Ewers (BPBM); 3 males, 1 female, Eio Creek, nr. Bamanumu, CL 1840, 22 September 1983, D. A. and J. T. Polhemus (JTPC). East Sepik Prov.: 1 male, Suanimbu, E. of Maprik, 180 m., 14 January 1960, T. C. Maa (BPBM); 1 female, Bainyik, nr., Maprik, 225 m., 20-21 July 1961, J. L. and M. Gressitt (BPBM). Eastern Highlands Prov.: 1 male, Kassam Pass, 1460 m., 18 August 1963, J. Sedlacek (BPBM); 1 male, 18 km. S. of Okapa, 1300 m., 2 June 1967, on water trickle, G. A. Samuelson (BPBM). Gulf Prov.: 2 males, 2 females, Lubu River above Omo, 50 m., water temp. 19°C., 1 March 1995, D. A. Polhemus (BPBM); 1 female, Wamivei Creek, near Veim Creek, Kikori River delta, SW of Kikori, 40 m., water temp. 25°C., 5 March 1995, D. A. Polhemus (BPBM). Madang Prov.: 1 male, 1 female, Seleo, Berlinhaf. [Berlinhafen], 1896, Biro (JTPC); 1 female, Adelbert Mtns., Wanuma, 800-1000 m., 26 October 1958, J. L. Gressitt (BPBM). Milne Bay Prov.: 3 males, 2 females, Modewa Bay, Modewa, 0-50 m., 22 December 1956, L. J. Brass, Fifth Archbold Expedition to New Guinea (JTPC); 1 female. Woodlark Is., Murua, Kulumadau Hill, 16 February 1957, W. W. Brandt (BPBM); 9 males, 4 females, Normanby Is., Wakaiuna, Sewa Bay, 11-20 December 1956, W. W. Brandt (BPBM); 1 male, 3 females, Naura, 5 km. W. of Milne Bay, 7 November 1965, R. Straatman (BPBM); 1 female, Mt. Dayman, Ma- neau Range, 700 m., N. slope, 13-20 July 1953, G. M. Tate (JTPC); 1 male, 1 female, S. of Alotau, 29 September 1989, R. N. B. Prior (OXUM). Morobe Prov.: 2 males, 4 females, Garaina, 800 m., 15—16 January 1968, J. and M. Sedlacek (BPBM); 1 female, 16 km. W. of Mumeng, 80 km. N. of Wau, 900-1500 m.. May 1962, J. Sedlacek (BPBM); 4 males, 7 females, Huon Peninsula, Finschhafen, 80- 200 m., 13 April 1963, J. Sedlacek (BPBM); 1 male, Huon Peninsula, Pindiu, 870- 1500 m., 21-22 April 1963, J. Sedlacek (BPBM); 1 male, Ulap, 800-1100 m., Sep- tember 1968, N. L. H. Krauss (BPBM); 1 female, Lae, July 1944, F. E. Skinner (BPBM); 1 female, Wampit Vally, nr. Gurakor village, 950 m., 7 July 1957, D. E. Hardy (BPBM); 1 female, Mt. Lawson, N. Kakoro Gulf, 50-200 m., 16 March 1974, Gressitt and Reni (BPBM). New Ireland Prov.: 3 males, 8 females, Gilingil Plain, 2 m., 16 July 1956, J. L. Gressitt (BPBM); 2 males, ridge above “Camp Bishop”, 15 km. up Kait River, 250-750 m., 14 July 1956, J. L. Gressitt (BPBM); 1 male, 1 female, old German road W. of Karu Plateau, 40 km. N. of Namatanai, 9 December 1969, J. E. Tobler (JTPC). Southern Highlands Prov.: 1 male, slow trib. to Mubi River, SE of Kantobo, 380 m., water temp. 22°C., 8 March 1995, D. A. Polhemus (BPBM); 2 males, 4 females, Auwa Creek, nr. Kafka, NW of Kantobo, 520 m., water temp. 22°C, 9 March 1995, D. A. Polhemus (BPBM); 2 males, 2 females, swift clear trib. to Digimu River, 1 km. above Sorotage, 400 m., water temp. 23.5°C., 10 March 1995, D. A. Polhemus (BPBM); 3 males, 7 females, Ofake Creek, trib. to Mubi River, 2 km. NW of Kantobo, 380 m., water temp. 23.5°C., 11 March 1995, D. A. 1997 LIMNOMETRA OF NEW GUINEA 35 Polhemus (BPBM); 1 male, 4 females, branch of upper Kara Creek (trib. to Digimu River), 3.5 km. S. of Moro oil camp, on road to Ridge camp, 900 m., water temp. 20°C., 13 March 1995, D. A. Polhemus (BPBM); 3 males, 2 females, small forest tributary to upper Mubi River at Swinging Bridge, nr. Tubage, NE of Moro, 900 m., water temp. 20°C., 14 March 1995, D. A. Polhemus (BPBM); 2 males, 1 female, small rocky creek in disturbed forest, approx. 1.0 km. N. of Tubage, NE of Moro, 1000 m., water temp. 21°C., 14 March 1995, 14:30-15:00 hrs.; 22 March 1995, D. A. Polhemus (BPBM); 1 male, impended roadside stream in forest at Moro oil camp, 840 m., water temp. 25°C., 14 March 1995, 16:00-17:00 hrs.; 24 March 1995, D. A. Polhemus (BPBM); 7 males, 5 females, small rocky stream on N. slope of Mt. Bosavi, 1250 m., water temp. 19°C., 16 March 1995, 14:30-17:00 hrs.; 17 March 1995, D. A. Polhemus (BPBM); 3 males, 2 females, swift rocky stream in moss forest on N. slope of Mt. Bosavi, 1400 m., water temp. 18°C., 19 March 1995, D. A. Polhemus (BPBM). East New Britain Prov.: 1 male. Gazelle Peninsula, upper Warangoi, 250-600 m., 28-30 November 1962, J. Sedlacek (BPBM). West Sepik Prov.: 1 male. Amok [nr. Aitape], 165 m., 6 January 1960, T. C. Maa (BPBM); 1 male, 3 females, Dreikkir, 350 m., 23 June 1961, J. L. and M. Gressitt (BPBM); 3 males, Telefomin, March-June 1971, A. B. Mirza (BPBM); 2 females, Torricelli Mtns., Mobitei, 750 m., 1-15 April 1959, W. W. Brandt (BPBM). Western Prov.: 1 male. Fly River, Olsobip, 400-500 m., August 1969, J. and M. Sedlacek (BPBM); 7 males, 3 females. Fly River above Kiunga, CL 1774, 3 September 1983, D. A. and J. T. Polhemus (JTPC). Western Highlands Prov.: 1 male, Baiyer River, 1150 m., 17 October 1958, J. L. Gressitt (BPBM); 1 male, Kumur, upper Jimi Valley, 1000 m., 13 July 1955, J. L. Gressitt (BPBM). INDONESIA: Irian Jaya Prov.: 1 female, Humboldt Bay Dist., Pukusam Dist., W. of Tami River, June 1937 (JTPC); 7 males, 4 females, Bernhard Camp, 100 m., 13 April 1939, L. J. Toxopeus, Neth. Ind.-American New Guinea Exp. (RMNH); 2 males, 5 females, Vogelkop, Bomberi, 700-900 m., 6 June 1959, T. C. Maa (BPBM); 3 males, 3 females, Biak Is., Kampong Landbouw, 50-100 m., 29 May 1959, J. L. Gressitt (BPBM); 1 male, Nabire, S. Geelvink Bay, 0-30 m., 2-9 July 1962, J. L. Gressitt (BPBM); 5 males, 3 females, Waris, S. of Hollandia, 450-500 m., 24-31 July 1959, T. C. Maa (BPBM); 3 males, Genjam, 40 km. W. of Hollandia, 100-200 m., 1-10 March 1960, T. C. Maa (BPBM); 1 male, 3 females, Kebar Valley, W. of Manokwari, 550 m., 4-31 January 1962, S. Quate (BPBM); 1 female, Hollandia (Kota Bharu), 13 March 1960, T. C. Maa (BPBM); 3 males, 2 females. Central Mtns., Archbold Lake, 760 m., 26 November-3 December 1961, S. and L. Quate (BPBM); 4 males, 7 females, W. of Sentani, 90+ m., 24 January 1959, T. C. Maa (BPBM); 3 males, 4 females, Fak Fak, S. coast Bomberai, 9 June 1959, T. C. Maa (BPBM); 2 males, 1 female. Star Mtns., Sibil Valley, 1245 m., 31 October 1961, S. Quate (BPBM); 12 males, 7 females, Salawati Is., Wajaar River, Wagom Mountains, W. of Sorong, CL 2623, 0-30 m. (0-100 ft.), water temp. 28°C., 30 Sept. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 2 males, 5 females, small forest trib. to Klagalo River at old Klagagi oil field, SE of Sorong, CL 2628, 45 m. (150 ft.), water temp. 25.5°C., 1 Oct. 1991, D. A. Polhemus (JTPC, BPBM); 4 males, 4 females, pools in muddy lowland rain forest streambed, 28 km. NE of Kasim on pipeline road, CL 2630, 15 m. (50 ft.), 2 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 5 males, 5 females, 8 immatures, upper Kali Mati River, 10 km. N. of Fak Fak, CL 2633, 36 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) 275 m. (900 ft.), water temp. 23°C., 10 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 1 male, pools in dry bed of middle Kali Mati River, 8 km. N. of Fak Fak, CL 2634, 230 m. (750 ft.), 10 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 8 males, 7 females, Mupi River, above Warkomi, Arfak Mountains, 38 km. S. of Manokwari, Cl 2646, 150 m. (500 ft.), water temp. 19°C., 18 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 7 males, 5 females, Saumarin River, nr. Warkomi, Arfak Mountains, 42 km. S. of Manokwari, CL 2647, 90 m. (300 ft.), water temp. 25°C., 18 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 4 males, 2 females, Kabori River, nr. Warkomi, Arfak Mountains, 41 km. S. of Manokwari, CL 2648, 15 m. (50 ft.), water temp. 25°C., 18 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 3 males, 3 females, small rocky stream at Aimasi Hulu, Arfak Mountains, 65 km. SW of Manokwari, CL 2649, 140 m. (450 ft.), water temp. 25°C., 19 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 7 males, 3 females, intermittent stream with pools at Kuala Kencana light industrial park, N. of Timika, 4°26.21 S, 136°51.84 E, 300 ft., water temp. 25°C., 17 January 1997, 14:00-15:00 hrs, and 20 January 1997, 09:30-11:30 hrs., CL 7043, D. A. Polhemus (USNM, LIPI); 4 males, 4 females, small forest streams and pools along Kopi River road, NE of Timika, 4°25.25 S, 136°56.44 E, 400 ft., water temp. 27°C., 18 January 1997, 08:00-11:00 hrs., CL 7044, D. A. Polhemus (USNM, LIPI); 4 females, small rocky stream at Mile 50 storage tank site on Tembagapura Road, 34 km. N. of Timika, 4° 16.99 S, 137°00.86 E, 2000 ft., water temp. 22°C., 19 January 1997, 09:00-10:00 hrs., CL 7046, D. A. Polhemus (USNM, LIPI); 1 male, pond and blackwater stream 2.4 km. S. of Mile 50 storage tank site on Tembagapura Road, 31.5 km. N. of Timika, 4°17.93 S, 136°59.98 E, 1900 ft., water temp. 25°C., 19 January 1997, 11:15-12:00 hrs., CL 7047, D. A. Polhemus (USNM); 1 male, 5 females, sago swamp and swift roadside drainage ditch at Km. 22 on Portsite road, 15 km. S. of Timika, 4°38.74 S, 136°53.93 E, 50 ft., water temp. 27°C., pH 7.45, 21 January 1997, 11:00-11:30 hrs., CL 7054, D. A. Polhemus (USNM, LIPI); 4 males, 5 females, upper Minajerwi River and swift tributary, approx. 1 km. E. of Mile 50 tank farm on Tembagapura Road, 4° 16.99 S, 137°01.56 E, 1650 ft., water temp. 23°C., 24 January 1997, 12:00-14:00 hrs., CL 7061, D. A. Polhemus (USNM, LIPI); 3 males, 3 females, small rocky stream along N. side of P. T. Freeport Indonesia Etna Bay exploration camp, head of Etna Bay, 0-60 m. (0-200 ft.), 3°58.10'S, 134°57.68'E, water temp. 25-27°C., 28-29 March 1997, CL 7077, D. A. and J. T. Polhemus (USNM). Maluku Prov.: 2 males, 1 female, Aru Archipelago, Trangan Is., 1 km. S. of Popjetur, 6°48'S, 134°4'E, 10 July 1994, A. H. Kirk-Spriggs (NMWC). Limnometra lipovskii (Kirkaldy) Figs. 10, 15 Limnometra lipovskii Hungerford & Matsuda 1958, Univ. Kansas Sci. Bull. 39: 399 (Type, macropterous male, Guadalcanal, SEMK). Discussion: Limnometra lipovskii is rather uncommon and localized in New Guinea, being found only in the coastal lowlands or along larger rivers (Fig. 15), and few series from the island are present in major collections. This species was overlooked completely by Nieser and Chen (1992) in the review of Limnometra of the Indo- 1997 LIMNOMETRA OF NEW GUINEA 37 Fig. 15. Distribution of Limnometra lipovskii Hungerford and Matsuda in New Guinea. Stippling indicates areas above 2000 m. Australian region; neither it or any other of the species occurring in New Guinea is included in their key. Distribution: New Guinea (Fig. 15), Solomons (type locality: Guadalcanal), Aus- tralia, Halmahera. New Guinea records: PAPUA NEW GUINEA: Gulf Prov.: 2 males, 2 females, roadside ponds in low- land forest, 2 km. S. of Kopi oil camp, N. of Kikori, 20 m., water temp. 24°C., 27 February 1995, D. A. Polhemus (BPBM). Western Prov.: 1 male, 1 female. Fly River above Kiunga, CL 1774, 3 September 1983, D. A. and J. T. Polhemus (JTPC); 1 male, 1 female. Fly River at Kiunga, CL 1864, 5 September 1983, D. A. and J. T. Polhemus (JTPC). East Sepik Prov.: 1 female, Ramumba, NW of Wewak, CL 1801, 11 September 1983, D. A. and J. T. Polhemus (JTPC). INDONESIA, Irian Jaya Prov.: 2 males, 1 immature, Biak Is., Kampong Land- bouw, 50-100 m., 27 May 1959, J. L. Gressitt (BPBM); 2 males, 3 females, swamp forest pond S. of Walio oil field, nr. Kasim, CL 2620, 7 m. (20 ft.), water temp. 30°C., 29 Sept. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 2 males, 1 female, Salawati Is., Wajaar River, Wagom Mountains, W. of Sorong, CL 2623, 0-30 m. (0- 100 ft.), water temp. 28°C., 30 Sept. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 10 males, 18 females, Batuputih River nr. Krooy, 3 km. NW of Kaimana, CL 2639, 30 m. (100 ft.), water temp. 26°C., 12 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 1 male, Airtiba River, 3 km. NW of Krooy, NW of Kaimana, CL 2640, 15 m. (50 ft.), water temp. 26°C., 13 Oct. 1991, J. T. and D. A. Polhemus (JTPC, BPBM); 1 male, wetlands in former Ajkwa River channel near Km. 21 on Portsite road, 16 km. S. of Timika, 4°39.91 S, 136°53.83 E, 50 ft., water temp. 26.5°C., pH 7.4, 21 January 1997, 10:00-11:00 hrs., CL 7053, D. A. Polhemus (NMNH); 3 males, 2 females, pond and blackwater stream 2.4 km. S. of Mile 50 storage tank site on Tembagapura Road, 31.5 km. N. of Timika, 580 m., 4°17.93 S, 136°59.98 38 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) E., water temp. 25°C., 5 March 1997, 11:00-12:00 hrs., CL 7047, D. A. and J. T. Polhemus (USNM, JTPC). Maluku Prov.: 1 male, 1 female. Am Archipelago, Tran- gan Is., 1 km. E. of Ngalgull, 90 m., 6°48'S, 134°4'E, 29 July 1994, A. H. Kirk- Spriggs (NMWC). ACKNOWLEDGMENTS We wish to thank the World Wildlife Fund and the Papua New Guinea Department of En- vironment and Conservation (DEC), who organized the Field Survey of Biodiversity in the Kikori River basin in early 1995 that led to the initial discovery of Limnometra grallator. Special thanks are also due to the following persons who aided us with field surveys in Papua New Guinea during 1983: David and Dorothy Softe, Balimo; Vance and Patty Woodward, Kiunga; David Coates and Moses Arumba, Wewak; Stan and Kris Niemi, Mt. Hagen; and especially John Ismay, formerly of the Bureau of Primary Industry, Konedobu. We are also pleased to acknowledge the invaluable assistance of the following persons who helped us with permits, guidance, transportation, and logistics in Indonesian New Guinea: Drs. Rosichon Ubaidillah, Dr. Soetikno Wiroatmodjo, Dr. Mohammed Amir, and other personnel from the Indonesian Institute of Sciences (LIPI), Bogor; Mrs. Moertini, Mr. Sanchoyo, the other staff members of the LIPI office in Jakarta; Mr. Benny Lesomar, Natrabu Corp., Biak; Mr. Lewis Zeigler, Mr. Jan Beiers, Mr. T. Loebis, Mr. Franz Petrusz and many others from the Petromer Trend oil company, Jakarta and Sorong; Mr. Neal Blackburn, Mobil Oil Indonesia Inc., Jakarta; Mr. James Carpenter, Mobil Oil Indonesia Inc., Fakfak; Mr. Petrus Manggai, Fakfak; Mr. Jacob Bakabessy and Mr. Bernardus Sambery, PHPA, Manokwari. We also wish to provide special acknowledgement to Howard Lewis and Kent Hortle of P. T. Freeport In- donesia, for facilitating surveys in the vicinity of Timika. The following curators kindly provided access to specimens held under their care (collection abbreviations are those used in the text): Dr. Robert Brooks, Snow Entomological Museum, Lawrence (SEMK); Drs. Richard Froeshner and Karl Krombein, National Museum of Natural History, Smithsonian Institution, Washington, D. C. (USNM); Dr. N. M. Andersen, Universitets Zoologisk Museum Copenhagen (ZMUC); Dr. G. Cassis, Australian Museum, Sydney (AMSA); Dr. H. E. Evans, (formerly at) Museum of Comparative Zoology, Harvard University, Cam- bridge (MCZC); T Weir, Australian National Insect Collection, Canberra (ANIC); Gordon Nish- ida. Bishop Museum, Honolulu; W Dolling, The Natural History Museum, London (BMNH); A. H. Kirk-Spriggs, National Museum of Wales, Cardiff (NMWC); Dr. Nico Nieser, Tiel, The Netherlands. Other collection abbreviations employed in the synonymies are as follows: ISNB = Institut Royal des Sciences Naturelles de Belgique; NHMW = Naturhistorisches Museum Wien; OXUM = University Museum, Oxford. The holotype of Limnometra grallator is deposited in the Bishop Museum, Honolulu (BPBM); paratypes are held in that collection, the J. T. Polhemus collection, Englewood, Col- orado (JTPC), and the Museum Zoologicum Bogoriense, Bogor (LIPI). This research was sponsored by a series of grants from the National Geographic Society, Washington, D. C. (2698-83, 3053-85, 3398-86, 4537-91), and by grant BSR-9020442 from the National Science Foundation, Washington, D. C. In addition, JTP conducted this research as an adjunct faculty member in the Dept, of Entomology at Colorado State University. We thank all these organizations for their continued support of our research into the systematics and zoogeography of aquatic Heteroptera. LITERATURE CITED Andersen, N. M. and T. A. Weir. 1997. The gerrine water striders of Australia (Hemiptera: Gerridae): taxonomy, distribution and ecology. Invertebrate Taxon. 1 1(2);203-299. 1997 LIMNOMETRA OF NEW GUINEA 39 Hungerford, H. B. and R. Matsuda. 1958. The Tenagogonus-Limnometra complex of the Ger- ridae. Univ. Kansas Sci. Bull. 39:371-457. Nieser, N. and R Chen. 1992. Revision of Limnometra Mayr (Gerridae) in the Malay Archi- pelago. Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), II. Tijds. Ento- mol. 135:11-26. Received 18 March 1997; accepted August 1997. J. New York Entomol. Soc. 105( l-2):40-44, 1997 A NEW SPECIES OF LEAF-MINING OULEMA FROM PANAMA (COLEOPTERA: CHRYSOMELIDAE; CRIOCERINAE) Fredric V. Vencl and Annette Aiello Department of Neurobiology and Behavior, State University of New York at Stony Brook, Stony Brook, New York 11794, and The Smithsonian Tropical Research Institute, Box 2072, Balboa, Ancon, Republic of Panama Abstract. — A new species of leaf-miner, Oulema pumila (Chrysomelidae:Criocerinae), from central Panama is described and illustrated. Host plant and life history data are given. One of the smallest species, it is only the second record of the leaf-mining habit and the first host record from the Piperaceae recorded for the subfamily. Leaf-mining appears to be derived in this instance and we offer some observations on its evolution. In 1994, one of us (A. A.) reared three beetles from mines in the leaves of an unidentified cultivated species of Peperomia (Piperaceae) in Arraijan, Panama Prov- ince, Panama. The new beetle belongs to the genus Oulema Des Gozis (Coleoptera: Chrysomelidae) and is one of the smallest members of the shining leaf beetle sub- family Criocerinae. The Criocerinae are a cosmopolitan group of nearly 2000 species noted for their smooth, often colorful appearance, narrow, unmargined pronotum and excrement-covered, folivorous larvae. Our report of leaf-mining larvae for this spe- cies is only the second one for the Criocerinae. Moreover, it is the first shining leaf beetle recorded from the Piperaceae. Although there is no universal agreement as to whether Oulema is a genus in its own right, or merely a subgenus within Lema (see for example Mohr, 1985), our use of Oulema follows the concept of White (1993) and Monros (1960). Members of the genus Oulema have the following distinguishing characteristics: antennal tu- bercles very close together or touching; angle of grooves separating vertex from the frons less than 90°; the vertex is often produced into tubercles; pronotal constriction weak, sub-medial to basal; antennae length longer than Vi the body length; elytra without color patterning; 9‘^ stria frequently interrupted for 2-8 punctures; aedeagus with lateral lobes more or less parallel and not concealing the medial lobe. Measurements were made with an ocular micrometer. Dimensions were taken from prominent morphological features: elytral length from humerus to apex, and total length from vertex to elytral apex along the meson because the head is directed downward. The habitus drawing was made using india ink, chalk, negro and charcoal pencils on Coquille board. Oulema pumila, new species Fig. 1 Diagnosis: Head and pronotum narrower than the elytra, claws connate. Extremely small size of 2.6 to 2.9 mm total length. The head, pronotum and abdomen are 1997 NEW O ULEMA FROM PANAMA 41 Fig. 1. Oulema pumila, dorsal habitus, holotype female. 42 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) yellowish orange. The head, is distinctly bituberculate at the vertex. The elytra are black with an iridescent violet tint. Description: Form sub-cylindrical. Total length 2.6 to 2.9 mm including the tuber- cles of the vertex. Head: yellowish orange, shiny, much narrower than the anterior width of the pronotum. Clypeus, labrum and mouth parts brownish black. Frons separated from the vertex by deep grooves that form an “X” with the angle between them less than 90°, the antennal tubercles almost touching and much closer together than the frons-vertex junction. Vertex deeply divided by a broad, longitudinal groove and produced laterally into two elliptical tubercles. Antennae dark brown, longer than Vi the length of the body, eleven segmented, filiform but widening very grad- ually toward the apex. Eyes very weakly notched by less than their width; head moderately constricted behind the eyes. Pronotum: Pale orange with a yellowish cast, shiny and emarginate; slightly wider than long; the apex slightly wider than the basal width. A shallow transverse constriction, 94% of the apical width, divides the pronotum sub-medially at its basal Vs and bears a large medial pit. Disc with two longitudinal, medial rows of 5-6 medium-sized pits. Elytra: Violet-black and shiny with an iridescent reflection; a transverse depression at the basal Vs; punctation of the 10 striae medium in size and widely spaced but well aligned; the 9'*^ stria incom- plete for 4-5 punctures; scutellum orange. Ventral side: Pale orange with brownish melanism, shiny and covered with sparse setae; punctation of mesostemum coarse, sparse and most dense near anterior edges; coxae pale yellowish orange. Legs: en- tirely light yellow with a brownish cast to the dorsal surfaces; two apical tarsal spurs; claws brownish-black, connate and simple. Holotype. Female: PANAMA, Panama province, Arraijan, 24 Sept. 1994. A. Aiello. Location: United States National Museum. Paratypes. 1 female and 1 male, same data as holotype. The female is deposited in the United States National Museum. The male is in the senior author’s collection. Variation. Paratype female 2.8 mm in length; paratype male 2.65 mm long with a brownish cast to ventral surfaces and the 9'*' stria interrupted for only 3 punctures. Pupation. 10 Sept. 1994. Eclosion. 24-25 Sept. 1994. Pupation Chamber. White, foamy in texture, 3.5 mm long, 2.1 mm wide. Host Plant. Peperomia sp. (Piperaceae). Etymology. From the Latin meaning dwarfish. Remarks: Although the aedeagus of the single male at our disposal was lost during dissection, it was observed to have had the sides of the lateral lobes parallel and not concealing the medial lobe, which conforms to the aedeagal characteristics for the genus Oulema. Oulema pumila is most closely allied to Lacordair’s (1845) second division that includes Lema species, sensu lato, that are distinguished by having red bodies and blue elytra with the 9'^ stria interrupted. Among the smallest species in the division, Oulema pumila can be differentiated from: L. tenella, by lacking a black abdomen; L. jocosa and L. concinna by lacking elytral markings of any kind; L. vidua and L. impura by lacking yellow elytra; L. stolida by lacking an entirely black body; and from L. gilveola by having a notched eye. Discussion: This tiny species is a rare denizen of Caribbean rain forest where it is a miner in the leaves of epiphytic pipers. This is the first host record for the Cri- 1997 NEW O ULEMA FROM PANAMA 43 ocerinae from the Piperaceae. Examination of herbarium material revealed Peper- omia macrostachya specimens with mines large enough to have been made by an insect the size of O. pumila. In addition, one pupal chamber in the leaf mesophyl was lined with a white, foamy substance typical of the buccal material produced by members of the subfamily Criocerinae to form pupation chambers. Monros (1960) mentions Lema quadrivittata de Borre as a leaf-miner of plants in the Commelinaceae of South America. Oulema pumila can be differentiated readily from L. quadrivittata by (1) the lack of elytral stripes, (2) size smaller than 4.2 mm and, (3) disparate host association. The genus Oulema as proposed by Des Gozis (1886), embraces New and Old World Lemiini that fall into the red body/blue elytra category. This cosmopolitan genus of leaf beetles, recently augmented by White (1993), now includes 20 North American species, of which host plants are known for 15. It is interesting to note that the larvae of Old World species feed on the Poaceae while their New World cousins feed mainly on members of the dayflower family Commelinaceae. This may well indicate that a very old split in the genus occurred well before the appearance of the grasses, perhaps as a consequence of the break-up of Gondwana some 135 million years ago. The total number of species assigned to the genus undoubtedly will increase following much needed revision and reassignment of Central and South American species. The addition of O. pumila brings the number of Panamanian species in the subfamily Criocerinae to 34 (Vend, in prep.). Larvae of the entire subfamily Criocerinae are noteworthy for their peculiar habit of accumulating excrement on their backs to form fecal shields. Recently, fecal shields were demonstrated to contain predator deterrent chemicals derived entirely from host-plant metabolites (Morton and Vend, 1998). When forced to feed on the surface, stem boring and probably leaf mining larvae form fecal shields (Richardson, 1893; Kaufmann, 1967; Vend, pers. ob.). The retention of the shield-forming poten- tial suggests that mining and boring are derived behaviors. Internal feeding modes obviate the need for a shield defense. When the proposed sister taxa to the Chrys- omelidae, the Bruchidae and Cerambycidae, all with internally feeding larvae, are considered together with the numerous instances of gallicoly, stem, root and leaf- mining larvae within the Chrysomelidae itself, it becomes evident that surface fo- livory by leaf beetle larvae is a derived mode of feeding (Crowson,1981). We suspect this to be the case because folivorous larvae, which are physically exposed, soft bodied and flightless, are more vulnerable to predators and parasitoids than are eggs, pupae or adults. These observations implicate natural enemies as major determiners of feeding behavior in leaf beetles in general and perhaps of leaf-mining in O. pumila in particular. Endophytism, which is uncommon in related criocerines, is a specific adaptation by O. pumila to the threat of predation and may well represent an evo- lutionary reversal. However, we cannot exclude the possibility that mining in this instance is a retained, ancestral habit until the host is analyzed for the presence or absence of compounds known to be effective in the shield defenses of Oulema pumila’ s relatives. LITERATURE CITED Crowson, R. A. 1981. The Biology of the Coleoptera. Academic Press, London. Des Gozis, M. 1886. Recherche de I’espece typique de quelques anciens genres. Monlucon: 5-36. 44 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Kaufmann, D. L. 1967. Notes on the biology of three species of Lema (Coleoptera: Chryso- melidae) with larval descriptions and a key to described species. J. Kansas Entomol. Soc. 40:361-372. Lacordaire, J. T. 1885. Monographic des Coleopteres subpentameres de la famille des Phyto- phages. 1(1). Mem. Soc. R. Sci. Liege 3:1-740. Mohr, K. H. 1985. Beitrage zur Insektenfauna der DDR: Coleoptera-Chrysomelidae: Donaci- inae, Orsodacninae. Beitr. Entomol. 35:219-262. Monros, E 1960. Los Generos de Chrysomelidae (Coleoptera). Opera Lilloana III. Univ. Nac. Tucuman: 1-337. Morton, T. and E Vend. 1998. Larval Beetles Form a Defense from Recycled Host Plant Chemicals Discharged as Fecal Wastes. J. Cem. Ecology: in press. Richardson, W. D. 1893. Notes on Lema sayi. Proc. Entomol. Soc. Wash. 2: 240. Vend, F. V. in prep. Observations on the taxonomy and host plant associations of shining leaf beetles (Chrysomelidae:Criocerinae) of Costa Rica and Panama. White, R. E. 1993. A revision of the Subfamily Criocerinae (Chrysomelidae) of North America North of Mexico. U. S. D. A. Technical Bulletin 1805:1-158. Received 28 November 1996; accepted 21 June 1997. J. New York Entomol. Soc. 105(1— 2):45— 49, 1997 NECTARINELLA XAVANTINENSIS, A NEW NEOTROPICAL SOCIAL WASP (HYMENOPTERA: VESPIDAE; POLISTINAE) SiDNEi Mateus and Fernando B. Noll Faculdade de Filosofia, Ciencias e Letras de Ribeirao Preto, Depto. de Biologia. USP. Av. Bandeirantes 3900. 14040-901 - Ribeirao Preto - SP - Brasil, e-mail: sidmateu@usp.br. fax: 55 016 6335015 Abstract. — Nectarinella xavantinensis, a new epiponine species from the Neotropics, is de- scribed and the nest illustrated. This is the second known species in the genus and the first described from South America. A comparison with the other species from this genus is made. Comments regarding the biogeographical distribution of the genus are added. Nectarinella Bequaert is a monotypic genus of social wasps. It belongs to the tribe Epiponini (Carpenter, 1993) of the family Vespidae, composed of polygynous, swarm-founding wasps. The only previously known species, Nectarinella championi (Dover, 1924) is recorded from Central America (Richards, 1978) and Colombia (Schremmer, 1977) and presents astelocyttarous nests (Richards, 1978; Wenzel, 1991). This paper describes a new species, Nectarinella xavantinensis, found in Mato Grosso, center-western Brazil. Nectarinella xavantinensis, new species Diagnosis: Predominantly brownish. Malar area reduced and bare, epistomal suture sinuate with rounded comers (Fig. Aa), clypeus wider than high (Fig. A). Head with sparse punctures with erect hairs, the interantennal area elevated forming a V-shaped protuberance (Fig. Bb). Pronotum grossly punctured with evident pronotal lip, acute pronotal lobule (Fig. Cc). Mesonotum and scutellum grossly punctured, there is a small space between the epistomal suture and the inner orbital margin in the lower part of the eye (Fig. Af). Description: Female: Mean forewing length 5.85 mm. Structure: frons sparsely punctured, with hairs in each minute puncture, covered with short pubescence. Clypeus wider than high and sinuous, usually with two short longitudinal indefinitely shaped markings, delimited by the epistomal suture (Fig. lAa). Epistomal suture with curved angles (Fig. lAa). Interantennal area forming an elevated protuberance (Fig. IBb). Malar area reduced and bare. Pronotum grossly punctured with hairs; dorsal pronotal carina evident, pronotal lobule forming an acute angle (Fig. ICc). Mesonotum with a re- duced carina. Scutellum with hairs arising from large punctures. Mesoscutum sparse- ly punctured with short hairs and two stripes extending longitudinally, wider at the edges (Fig. ICd). Post-scutellum slightly narrower than scutellum, with parallel sides, anterior margin transverse, posterior margin rounded, shiny and almost im- punctate; propodeum wider than long, broadly and shallowly depressed medially. 46 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Fig. 1. Head (A, B) and thorax (C) of Nectarinella xavantinensis. Other explanations in the text. more or less impunctate, with comparatively long hairs on the sides. The first two segments of the abdomen with recumbent hairs arising from fine punctures, giving a finely striated appearance; succeeding segments very closely punctate with short hairs. Legs with pubescent hairs maculae close to the articulations. Color — head black and yellow, covered with short golden pubescence, clypeus yel- low with hairs, usually with two brownish-light markings; interanntenal area yellow. 1997 NEW NECTARINELLA 47 genae yellow having an ochraceous area extending to the occiput just behind the middle of each posterior ocellus; paraocullar spot yellow, continuous, reaching the occiput, frons black with median line above the antennal sockets. Eyes dark brown; antennae mostly ochraceous, lighter on the underside, outer side of flagellum, except the three apical articles, brownish-dark. Mandibles light brown, pronotum ochraceous with short golden brown hairs arising from large punctures, yellow line at the su- perior margin of the dorsal pronotal carina. Mesonotum yellow-ochraceous, scutel- lum predominantly yellow with golden brown hairs. Mesoscutum predominantly black with golden short hairs and two curved stripes. Post-scutellum yellow, pro- podeum yellow with comparatively long golden brown hairs on the sides. Abdomen brownish, the first two segments with dark brown short hairs, suceeding segments with short golden hairs, third to sixth segments brownish-light with lateral margin yellow; seventh segment ochraceous. Legs ochraceous with yellow maculae close to articulations, wings iridescent-hyaline, with cubito-radial and costal area darker than the rest, veins brownish-light with short hairs, tegulae ochraceous. Male: Unknown. Type material: holotype female, Brazil: Novo Sao Joaquim (Mato Grosso - April 04, 1996), Nova Xavantina region (14°45'S, 52°55'W). Collected by Mateus, S. & Noll, EB. Holotype and two paratypes deposited in the collection of Museu de Zoologia — USP, Brazil; two paratypes in the Museu E. Goeldi, Brazil; two paratypes in the American Museum of Natural History. Etymology: the specific name N. xavantinensis is a reference to the brazilian town of Nova Xavantina in the state of Mato Grosso. Nest: Astelocyttarus. Found Apr. 04 1996, 80 km west of Nova Xavantina (center- western Brazil). The nest was located 3.5 meters high in a Psidium sp. tree (Myr- taceae), in a secondary branch with 30° slope in relation to the ground. Comb with 103 cells, all of them with eggs; no meconia were found, indicating that the nest was very young. Envelope: Made of vegetal fibers mixed with salivary substances, predominantly light gray; with longitudinal green stripes (Fig. 2). Nest entrance located in the lower portion of the envelope, ring-shaped and 5 mm in diameter. As in N. championi the bark around the envelope showed small droplets of sticky substance which may serve against ant protection (Schremmer, 1977). Comb: Located centrally, with lateral spaces between the envelope and cells, 2 cm wide above and 4 cm below. All cells were sessile, directly built on the substrate using the same material as the envelope. The cells were 3. 5-3. 7 mm wide. At the center of the comb, complete cells were 4 mm high. Lower cells usually 1 mm high were located at the periphery of the comb (Fig. 2). The cells had no paper bottom and the eggs were laid on the cell walls very close to the bottom. No droplets of stored nectar were found in any of the cells. Remarks: Nectarinella xavantinensis, presents several morphological differences distinguishing it from N. championi. The most conspicuous features separating N. xavantinensis from N. championi are as follows. Head: Nectarinella xavantinensis has sparses punctures with erect hairs. The inter- antennal area is more elevated in Nectarinella xavantinensis (Fig. IBb), forming a V-shaped protuberance. The paraocular spot reaches the occiput in Nectarinella xav- antinensis (Fig. lAe) but in Nectarinella championi it is interrupted in the occelli 48 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Eig. 2. Nest (comb — left and envelope — right) of Nectarinella xavantinensis. height. The clypeus is wider than high in Nectarinella xavantinensis (Fig. lA), in Nectarinella championi it is as wide as high. In Nectarinella xavantinensis epistomal suture is sinuate, with rounded comers, especially in the anterior tentorial fovea. There is a small space between the epistomal suture and the inner orbital margin in the lower part of the eye (Fig. lAf). In Nectarinella championi the space between the tentorial fovea and the epistomal suture forms a 90° angle. This suture reaches the mandible passing through the inner orbital margin. Pronotum: In Nectarinella xavantinensis grossly punctured with short hairs and the pronotal lip is much more evident than in Nectarinella championi. The pronotal lobule in Nectarinella xavantinensis (Fig. ICc) is acute and in Nectarinella cham- pioni is around. Mesonotum: grossly punctured with a small sulcus in the anterior part. In Nectari- nella championi this sulcus is more evident with iridescent hairs and short bristles. The scutellum in Nectarinella xavantinensis is grossly punctured with erect bristles. In Nectarinella championi the punctures are thiner. Abdomen: Nectarinella xavantinensis — Brownish with fine punctures, short hairs and erect bristles close the margins. First segment with central spot, second segment with marginal maculae and sparse hairs. Other segments with stripes of yellow spots near the margins, short hairs and erect bristles. In Nectarinella championi the abdomen is predominantly ochraceous with stripes at the distal margins, and short hairs. Legs: Nectarinella xavantinensis has maculae close to articulations and pubescent hairs. In Nectarinella championi these maculae are absent. Wings: Nectarinella xavantinensis — veins brownish-light with short hairs and the area between the cells Cu radial and costal are darker than the rest. In Nectarinella championi short hairs are spread in all wings regions. 1997 NEW NECTARINELLA 49 Finally, the biogeographic distribution of both Nectarinella species is remarkable especially because it differs from other groups with wide distribution such as Agelaia and Polybia (Richards, 1978). These two species are vicariant. However, their pres- ence in two disjunct areas reflects either scarce field collections or truly disjunct distribution. Evidently, the solution will depend on additional collecting. ACKNOWLEDGMENTS The authors acknowledge the financial support by Fapesp (Funda^ao de Amparo a Pesquisa do Estado de Sao Paulo), Department of Biology (FFCLRP-USP), Zilia P L. Simoes (President of the Post-graduation Committee, Entomology Program — FFCLRP-USP), Dalton S. Amorim and Ronaldo Zucchi for their reading through the manuscript. LITERATURE CITED Carpenter, J. M. 1993. Biogeographic patterns in the Vespidae (Hymenoptera): Two views of Africa and South America. In: Peter Goldbatt (ed.). Biological relationships between Africa and South America. Yale Univ. Press, New Haven and London. Dover, C. 1924. Notes on the genus Nectarina Shuckard (Vespidae). Psyche 31(6):305-307. Richards, O. W. 1978. The social wasps of the Americas excluding the Vespinae. British Museum (Natural History), London. Schremmer, F. 1977. Das Baumrinden-Nest der neotropischen Faltenwespe Nectarinella cham- pioni, umgeben von einem Leimring als Ameisen-Abwehr (Hymenoptera; Vespidae). Entomol. Germ. 3(4):344-355. Wenzel, J. W. 1991. Evolution of the nest architecture. In K. G. Ross and R. W. Matthews (eds.). The social biology of wasps. Cornell Univ. Press, Ithaca: 480-519. Received 22 October 1996; accepted 21 June 1997. J. New York Entomol. Soc. 105(l-2):50-64, 1997 DESCRIPTION OF IMMATURE STAGES OF TWO SPECIES OF PSEUDOLAMPSIS (COLEOPTERA: CHRYSOMELIDAE) AND THE ESTABLISHMENT OF A NEW COMBINATION IN THE GENUS Sonia A. Casari and Catherine N. Duckett Museu de Zoologia, Universidade de Sao Paulo, CP 42 694, 04299-970 Sao Paulo SP, Brasil; and Departamento de Biologia, Universidad de Puerto Rico, P O. Box 23360, San Juan, Puerto Rico 00931-3360 Abstract. — The taxonomic history of species of Pseudolampsis is discussed and a new com- bination, Pseudolampsis darwini is formed. Distigmoptera darwini which had previously been synonymized with P. guttata is shown to be a unique species based on characters of larval and genitalic morphology. The immature stages of Pseudolampsis guttata and the third instar larva and pupa of Psedolampsis darwini are described and figured. These larvae are compared with other known larvae of Alticini. Resumen. — Se descute la historia taxonomica del genero Pseudolampsis y se forma la nueva combinacion de Pseudolampsis darwini. Se demuestra que Distigmoptera darwini, previamente smomizada con P. guttata es especie unica, basandose en las caracteristicas de la larva y la morfologia de la genitalia. Se describen y trazan las etapas inmaduras de Pseudolampsis guttata y el ultimo estadio y la pupa de Pseudolampsis darwini. Estas larvas se comparan con otras larvas conocidas de Alticini. One of the easiest to recognize but most poorly known groups in the Alticini are the “monoplatines”, a group of small elongate beetles with continuous pubescence covering body and elytra, with very enlarged metafemora, globosely swollen last tarsal segment and nine elytral striae (Scherer, 1962). This is a neotropical group with 37 genera, over half described by Clark (1860) and first proposed as a group, as the Monoplatites, by Chapuis (1875). The monoplatines are not collected in large numbers and are poorly understood both taxonomically and biologically (Flowers and Tiffer, 1992). Despite the phenetic similarity of the monoplatine genera, the tribe Monoplatini (Leng, 1920) has not been universally accepted (Furth, 1988; Seeno and Wilcox, 1982) nor have any hypotheses of relatedness among the genera been pro- posed. Only one species of monoplatine larva has been described previously (Buck- ingham and Buckingham, 1981) and few species have corroborated records of host plant (Jolivet and Hawkeswood, 1995; Flowers and Janzen, 1997) or breeding habits (Flowers and Tiffer, 1992). The larvae of Alticini are in general poorly known, especially given the large size of the tribe (but see Lawson, 1991, for review). This may be due to life history of larvae, some larvae are nocturnal (Duckett, unpublished data), others are root or stem feeders (Lawson, 1991). However difficult larval data can be to obtain, larval morphology can be significant to the identification of closely related taxa as well as the resolution of higher level taxonomic relationships (Lawrence and Newton, 1995; Reid, 1995). Here we present a detailed study of larval morphology of Pseudolamp- sis, a monoplatine feeding on the waterfern, Azolla, to make these data available to 1997 PSEUDOLAMPSIS 51 coleopterists working at higher taxonomic levels, as well as to aide general under- standing of larval Chrysomelidae and of the monoplatines. We also present the tax- onomic history of genus Pseudolampsis and discuss the adult characters which sup- port the formulation of a new taxonomic combination in the genus. TAXONOMIC HISTORY Pseudolampsis was first described in 1889 by Horn, who transferred Hypolampsis guttata Leconte 1884 to Pseudolampsis. This genus was thought to be known only from the southern U.S. until Balsbaugh (1969) synonymized Distigmoptera darwini, known only from Uruguay and Mato Grosso, Brazil with P. guttata. Immatures of P. guttata from the U.S. were initially described by Buckingham and Buckingham (1981). Immatures of Pseudolampsis were collected in Sao Paulo State, Brazil, and as the original larval description (Buckingham and Buckingham, 1981) lacked detailed chaetotaxy and description of mouthparts, redescription was warranted. During the course of preparing the larval description (and study of the adult) it became apparent that the specimens collected in Sao Paulo (Fig. 1) represent a distinct species from those collected in Florida, USA. Specimens collected in Florida represent Pseudo- lampsis guttata (LeConte), a species known to feed on Azolla (Buckingham and Buckingham, 1981; Habeck, 1979). Specimens collected in Sao Paulo State proved to be conspecific with Distigmoptera darwini Scherer, 1964. Distigmoptera darwini is congeneric with Pseudolampsis, but the differences existing in the male and female genitalia as well as the larva support its individual identity. Balsbaugh figured the median lobe of the aedeagus of Distigmoptera darwini, collected in Mato Grosso, Brazil. This material obviously shows a groove in the ventral surface of the aedeagus, not present in P. guttata (cf. figs. 4 and 5 in Bals- baugh, 1969). This groove is present in a paratype of Distigmoptera darwini (which proved to be male on dissection despite Scherer’s assertion that all specimens were female (Scherer, 1964:298)), as well as in the Sao Paulo material. Dissection of the female genital system reveals a very similar system to that of Microdonacia (Reid, 1992: fig. 44). The bursa copulatrix is elongate, as are the vaginal palpi; the palpi are also presented in a unified pair apically (Figs. 2C-E), however, the basal area (or internal apodeme of Reid, 1992) is fused in Pseudo- lampsis and dorsally recurved. The eighth stemite (Fig. 2B) (spiculum gastrale, or tignum of Konstantinov (1994)) has an elongate basal portion, widens to a truncate setose apex; the epiproct and pygidum are also apically setose. In Pseudolampsis the spermathecae of both species are practically identical (see Fig. 2A); both possessing a flange on the pump, an enlarged proximal spermathecal duct, and a greatly enlarged gland valve. The vaginal palpi, however, are significantly different (Figs. 2C-E); in P. guttata (Fig. 2D, E) the internal apodeme is basally wider than the apex and in P. darwini it is narrower (Fig. 2C). Both possess 7 setae per palp. The bursa copulatrix in P. darwini is vested with microtrichea over its entire surface. In P. guttata only the opening of the bursa has microtrichea, which are significantly longer than those in P. darwini. The eighth stemite also differs between species in setation, which is much sparser in P. darwini. Differences in the larvae will be described and discussed below. These and the 52 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Fig. 1. Pseudolampsis darwini (Scherer), dorsal habitus, length 2.2 mm. 1997 PSEUDOLAMPSIS 53 Fig. 2. Pseudolampsis Female genitalia. A, spermatheca; B, eighth sternite ventral view, P. guttata-, C, vaginal palpi P. darwini (dorsal view); D-E, vaginal palpi P. guttata (dorsal, lateral). 54 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) mentioned characteristics of the genitalia warrant the establishment of a new com- bination. The formal synonomy is as follows; Pseudolampsis darwini n. comb. Distigmoptera darwini Scherer, 1964 Pseudolampsis guttata Balsbaugh, 1969 Pseudolampsis darwini (Scherer, 1964) Figs. 3A-H, 4A-J Larva (Figs. 3A,B) Length: 2-5 mm. General integument whitish; head brown with mandible yellowish-brown; anten- nae, maxillae and labium partially membranous; thorax and abdominal segments 1- 8 densely asperate, with sclerites bearing prominent setae and small pigmented spots; segments separated by grooves; spiracles annular located in darker sclerites. Flead rounded (Fig. 3C, D) moderately pigmented and sclerotized; frontal arms V- shaped; epicranial stem short; endocarina extending anteriad to epicranial stem, not reaching anterior margin. Frons bearing 3 pairs of hairy setae and 1 pair short scamiform setae; each epicranial half bearing 10 setae (7 dorsal, 3 ventrolateral). One convex, pigmented stemma each side. Antenna 2-segmented (Fig. 4E); membranous socket band- like, located at the end of frontal arms; basal segment partially membranous, bearing 2 dorsal sensoria on membranous area; distal segment cupuliform, sclerotized basally. Clypeus (Fig. 4A) transverse, narrow, sclerotized at basal half, bearing 2 setae on each side. Labrum (Fig. 4A) transverse, subtrapezoidal, slightly emarginate anteriorly, bearing 2 pairs of setae (lateral longer) and 1 pair of sensory pores. Epipharynx (Fig. 4B) densely covered by microtrichiae, concentrated in median anterior region; anterior margin bearing 6 pairs of stout pedunculate setae, 2 groups of campaniform sensilae near anterior margin and 2 groups at base on membranous area; 2 elongate darker areas near middle, each bearing a minute seta at apex. Mandibles (Figs. 4F, G) symmetrical, palmate, 5-toothed, dentae 2 and 3 serrate; external face bearing 2 setae and 2 sensory pores; penicillus formed by ramified setae. Maxilla (Figs. 4C, D); stipes elongate with 2 sclerotized areas, one small tranverse, near palp bearing 3 setae (1 short) and other basal larger, bearing laterally 2 ventral and 1 dorsolateral setae; cardo elongate, glabrous; mala bearing 6 moderately long pedunculate setae ventrally and 8 stout pedunculate dorsally (4 basal serrate and bunched); mala bearing microtrichiae dorsally; maxillary palp 3-segmented, 2 basal segments sclerotized at base; basal segment band-like bearing 1 ventral sensory pore; 2nd segment bearing 2 ventral and 1 dorsal setae; distal segment bearing ventrally 1 lateral sensory pore and dorsally 1 short seta and 1 sensillum placodeum. Labium (Figs. 4C, D): prementum membranous with a transverse sclerotized area bearing 2 setae; postmentum membranous bearing 1 well developed and 1 short seta and 1 sensory pore on each side and 2 minute setae near base; labial palp 2-segmented; basal segment with a ventrolateral sensory pore; distal segment with 1 ventrolateral sensory pore and 1 seta and 1 placoid sensillum dorsal. Hypopharynx (Fig. 4D) membranous, partially covered by microtrichiae; anteriorly bearing 6 setae (4 minute and 2 short); 2 longitudinal scle- rites. Gular area absent. Prothorax narrower than other thoracic segments; pronotum with sclerotized setose shield-like plate, divided at mid-line by whitish narrow band; each 1997 PSEUDOLAMPSIS 55 Fig. 3. Pseudolampsis darwini (Scherer). Larva: A, B, habitus (lateral, dorsal); C, D, head (dorsal, ventral); E, F, prothoracic leg (laterointernal, laterosternal). Pupa: G, H, dorsal, lateral. Figs. A, B; C-F; G, H, respectively to same scale. 56 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Eig. 4. Pseudolampsis darwini (Scherer). Larva: A, clypeus and labrum; B, epipharynx: C, D, maxillae and labium (ventral, dorsal); E, antenna (dorsal); E, G, mandible (internal, external); H, 6th tergite; I, 5th glandular opening; J, mesothoracic spiracle. Figs. A, C; E, I; E, G, re- spectively to same scale. 1997 PSEUDOLAMPSIS 57 side bearing 7 setae and minute darker spots scattered among them. Meso- and meta- thorax gradually wider than prothorax; both with a transverse median groove forming 2 plicae and identical arrangement of sclerites: 2 rounded sclerites on 1st plica, each bearing one seta, and 4 smaller sclerites on 2nd plica, each bearing 1 seta; each side with one dorsolateral sclerite bearing 2 setae; small sclerotized pigmented spots densely scattered among larger sclerites making irregular plates; intersegmental area between pro- and mesothorax with a lateral membranous prominence bearing an annular spiracle (Fig. 4J). Thoracic segments with 1 sclerite and 1 small lobe bearing 1 seta, lateral to each coxa. Legs (Figs. 3E-F) increasing in size from pro- to metathorax, 4-segmented setose and partially membranous; basal segment bearing 5 setae; 2nd segment bearing 10 setae; 3rd segment bearing 7 setae; tarsungulus bearing 1 seta and pul villus. Abdominal segments 1-7 divided dorsally by a transverse groove forming 2 plicae; segment 1 with 2 series of sclerites arranged into transverse rows: one with 4 and other with 6 rounded sclerites each bearing 1 seta; segments 2-5 with 2 rows of sclerites, each with a median larger sclerite bearing 2 setae (larger on first row) and 2 smaller on each side, each bearing 1 seta; segments 6-7 (Fig. 4H) with 1st plica similar to the preceding one and larger median sclerite of 2nd plica fused to one lateral on each side and bearing 4 setae; segment 8 with an irregular sclerite bearing 6 dorsal setae; segment 9 almost totally sclerotized dorsally, bearing 10 setae. Segments 1-8 with paired dor- solateral glandular opening (Fig. 41): first opening located anterior to the sclerite on first plica; 2-7 located posterior to rounded lateral sclerite on first plica, near groove; 8 near apex. Segments 1-8 with lateral paired annular spiracles located in sclerotized rounded lobe and paired lateral partially sclerotized lobes each bearing 2 setae. Ventrally, seg- ments 1-8 with 3 membranous lobes at middle (disposed in a triangle) each bearing 2 setae and 1 slightly sclerotized lobe, each bearing 3 setae. Pupa (Figs. 3G-H) Length: 2. 5-2. 8 mm Cream, bearing long brownish setae inserted in small tubercules. Head invisible from above, bearing 6 pairs of setae (2 pairs shorter). Prothorax bearing 7 pairs of dorsal setae and 1 pair of lateroposterior round spiracles; meso- and metanotum bearing 2 pairs of setae each; each femur bearing 1 pair of setae near apex. Abdom- inal segments 1-6 bearing 4 pairs of dorsal setae; segments 1-5 bearing a pair of laterodorsal round spiracles; segment 6 with a pair of vestigial rounded spiracles; segments 7-8 apparently bearing 2 pairs of short lateral setae; segment 9 with 2 distal projections, each bearing 2 short setae near base. Material examined. BRAZIL. Sdo Paulo: Guapiara, Fazenda Intervales (Sede de Pesquisa) (marsh), 09.xi.l992. (MZSP), 36 larvae, 7 pupae inside pupal cocoons and 4 adult fixed (MZSP). Larvae were prepared in glycerine. Pseudolampsis guttata (LeConte, 1884) Figs. 5A-G, 6A-N First instar larva (Fig. 5A) Length: 1.0-1. 5 mm General integument (including head) dorsally brownish, almost totally sclerotized, covered by setose sclerites very closely placed or fused; ventrally, membranous, whitish. 58 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Fig. 5. Pseudolampsis guttata (LeConte), Larva: A, first instar (lateral); B, C, mature (lat- eral, dorsal); D, antenna; E, F, prothoracic leg (laterosternal, laterointernal); G, head (dorsal). Figs. B, C; E-F, respectively to same scale. 1997 PSEUDOLAMPSIS 59 Head and mouth parts similar to mature larva. Stemmata large and translucent. Prothorax: pronotum with shield-like plate divided at middle by whitish narrow band, each side bearing 7 setae; each side with 1 lobe slightly sclerotized bearing 2 setae and 2 small sclerites each bearing 1 seta near coxa. Meso- and metathorax with a transverse median groove forming 2 plicae and an identical arrangement of sclerites: each plica with 1 dorsal transverse sclerotized plate; plate of first plica shorter, par- tially divided by reticulate area bearing 2 setae, the second, entire bearing 4 setae; each side with a rounded plate bearing 3 setae; 2 sclerites lateral to each coxa, the posterior bearing 1 seta; intersegmental area between pro- and mesothorax with a lateral sclerite bearing an annular spiracle. Legs similar to mature larva. Abdominal segments 1-7 divided dorsally by a transverse groove forming 2 plicae with an identical arrangement of sclerites: first plica with a median sclerite bearing 2 setae and 2 smaller sclerites on each side, each bearing 1 seta; 2nd plica with a larger median sclerite bearing 4 setae and 1 smaller seta on each side, each bearing 1 seta; these sclerites are very near each other, separated only by a small groove; segments 8-9 with a larger dorsal sclerite bearing respectively 2 and 6 setae. Seg- ments 1-8 with a paired dorsolateral glandular opening at same position as mature larva; each with lateral paired annular spiracles located in sclerotized rounded lobe; each segment with a lateral sclerotized lobe bearing 2 setae; ventrally with 3 mem- branous lobes at middle (arranged in a triangle) each bearing 2 setae and 1 larger sclerite each side, bearing 3 setae. Mature larva (Figs. 5B, C) Length 2. 5-4.0 mm General integument whitish; head brown with mandibles clearer; antennae, maxilla and labium partially membranous; thorax and abdominal segments 1-8 densely as- perate, with sclerites, bearing long setae and small pigmented spots giving dorsal integument a brown appearance; spiracles annular, located in darker sclerites. Head rounded (Fig. 5G), pigmented and sclerotized; frontal arms V-shaped; epi- cranial stem short; endocarina extending anteriad of epicranial stem, not reaching anterior margin; frons bearing 3 pairs of hairy setae and 1 pair of shorter and scam- iform setae medioanteriorly; each epicranial half bearing 10 setae (1 pair very short). One convex pigmented stemma on each side. Antenna 2-segmented (Fig. 5D); mem- branous socket band-like located at the end of frontal arms; basal segment partially membranous bearing 2 dorsal sensoria on membranous area and 2 sensory pores at border of sclerotized area; distal segment cupuliform, sclerotized basally. Clypeus (Fig. 6C) transverse, narrow, slightly sclerotized on basal half, bearing 2 short setae on each side. Labrum (Fig. 6C) transverse, slightly sclerotized, emarginate anteriorly bearing 2 pairs of setae (lateral longer) and 1 pair of sensory pores. Epipharynx (Fig. 6D) apparently partially covered by microtrichiae, more concentrated and lon- ger at median anterior region; anterior margin bearing 7 pairs of stout pedunculate setae (2 pairs near middle shorter; 1 pair bifurcate), 2 groups of sensillae near anterior margin, 2 groups near base and 2 larger sensillae near middle. Mandibles (Figs. 6G, H) symmetrical, palmate, 5-toothed; external face bearing 2 setae and 2 sensory pores; penicillus well developed, formed by ramified setae. Maxilla (Figs. 6A, B): stipes elongate with 2 sclerotized areas, one small, transverse, near palp bearing 3 setae, other area basal, longer, bearing 2 ventral and 1 dorsolateral setae; cardo elongate, glabrous; mala ventrally bearing 8 pedunculate setae, and dorsally partially 60 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Fig. 6. Pseudolampsis guttata (LeConte), Larva: A, B, maxilla and labium (ventral, dorsal); C, clypeus and labrum; D, epipharynx; E, 6th tergite; F, abdominal segment 2 (lateral) showing the glandular opening and spiracle; G, H, mandible (internal, external). Figs. A, B; C, G, H, respectively to same scale. 1997 PSEUDOLAMPSIS 61 covered by microtrichiae, and bearing 6 stout pedunculate setae with serrate apex; maxillary palp 3-segmented; 2 basal segments sclerotized at base; basal segment band-like, bearing 1 ventral sensory pore; 2nd segment bearing 2 ventral and 1 dorsal setae; distal segment bearing ventrally 1 lateral sensory pore and dorsally 1 seta and 1 sensillum placodeum. Labium (Figs. 6A, B): prementum with a transverse scler- otized area bearing 2 setae on anterior border and 2 sensory pores on posterior; postmentum membranous, each side bearing 1 long and 1 minute seta and 1 sensory pore, 2 minute setae near base; labial palp 2-segmented; basal segment with a ven- trolateral sensory pore; distal segment with a ventrolateral sensory pore and a dor- solateral placodeum sensillum. Hypopharynx (Fig. 6B) membranous, partially cov- ered by microtrichiae, bearing 6 minute setae; 2 longitudinal sclerites. Gular area absent. Prothorax narrower than other thoracic segments; pronotum with sclerotized se- taceous shield-like plate, divided at mid-line by whitish narrow irregular band, each side bearing 7 setae with minute darker spots scattered among them; one ventral lobe slightly sclerotized bearing 2 setae near coxa. Meso- and metathorax gradually wider than prothorax, both with a transverse median groove forming 2 plicae and an identical arrangement of sclerites: on first plica 2 small rounded sclerites, each bearing 1 long setae. On 2nd plica 2 sclerites of same size at middle, and 1 larger each side, each bearing one seta; each side with a large sclerotized tubercle each bearing three setae (1 shorter); small sclerotized pigmented spots densely scattered among larger sclerites; intersegmental area between pro- and mesothorax with a slightly sclerotized lobe bearing an annular spiracle. Each thoracic segment lateral to coxae with 1 sclerite and 1 lobe, each bearing 1 seta. Legs (Figs. 5 E, F) increasing in size from pro- to metathorax; 4-segmented, setaceous and partially membranous; basal segment bearing 5 setae; 2nd segment bearing 1 1 setae; 3rd segment bearing 6 setae; tarsungulus bearing 1 setae and pul villus. Abdominal segments 1-7 divided dorsally by a transverse groove forming two plicae; segment 1 with 4 sclerites on 1st plica, each bearing 1 seta, 2nd plica with 1 median larger sclerite bearing 4 setae and 1 smaller sclerite on each side, each bearing 1 seta; segments 2-7 (Fig. 6E) with 1 median larger sclerite bearing 2 setae and 2 smaller sclerite each side, each bearing 1 seta, and 2nd plica with 1 median larger sclerite bearing 4 setae and 1 lateral smaller sclerite each side, each bearing 1 setae, segment 8 with 1 dorsal irregular sclerite bearing 6 setae and 2 short setae, each side, on membranous area; segment 9 slightly sclerotized dorsally almost to- tally, bearing 10 setae; segment 10 membranous and ventral. Segments 1-8 with a paired dorsolateral glandular opening (Fig. 6F): first opening anteriorly to lateral sclerite of first plica; 2-7 between the 2 lateral sclerites on first plica near groove; 8 near apex. Segments 1-8 with lateral paired annular spiracles located in a rounded sclerite, and with paired lateral lobes (behind spiracle) each with sclerotized apex, each bearing 2 setae. Ventrally, segments 1-8 with 3 median small lobes disposed in a triangle, each bearing 2 setae and 1 larger lobe each side, each bearing 3 setae. Pupa Similar to P. darwini, differing only by presence of 5 pairs of setae on the head (6 in P. darwini). Material examined. USA. Florida. Alachu + Citrus Cos.: Gainesville and Crystal River, X. 1979, M. & G. Buckingham cols., 21 first instar and 35 mature larvae, 11 62 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) pupae fixed (MZSP) and 7 adults (2 MZSP and 5 collection C.N. Duckett). Larvae were prepared in glycerine for dissection. BIOLOGICAL NOTES Larvae and adults of Pseudolampsis darwini were collected on the Azolla sp. (Azollaceae) that was covering the pond surface. The pupae were found inside co- coons attached under or among the Azolla stems. The material of P. guttata was collected on Azolla caroliniana. Alticini larvae live on leaves or stems, on roots or underground parts of plants (Lawson, 1991). REMARKS Comparing the first instar with mature larva of Pseudolampsis guttata we verified that in the first instar the dorsal sclerites are larger or fused covering the dorsal integument almost entirely and the stemmata are larger and translucent, while in the mature larva the sclerites are smaller surrounded by small pigmented spots and the stemmata are pigmented. Other similarities are morphology of mouthparts, pronotum bearing 14 setae, abdominal segments 1-8 with a paired glandular opening and paired annular spiracles located in sclerotized lobes. Comparing the mature larva of Pseudolampsis darwini with P. guttata, they pres- ent different arrangment of sclerites of the abdominal segments (Figs. 3B, 4H; 5C, 6E, F) and different number of setae on anterior margin of epipharynx (Figs. 4B; 6D), mala (Figs 4C, D; 6A, B), and legs (Figs. 3E, F: 5E, F). Both species present similar frontal arms (Figs. 3C; 5G), and equal number of setae on pronotum (Figs. 3B; 5C), frons and epicranium (Figs. 3C; 5G), clypeus and labrum (Figs 4A; 6C), prementum (Figs 4C, 6A) and hypopharynx (Figs. 4D; 6B), and antennae 2-seg- mented (Figs. 4E; 5D), 1 pigmented stemma on each side (Figs. 3C; 5G), mandibles 5-toothed with penicillum ramified (Figs. 4F, G; 6G, H), maxillary palpi 3- and labial palpi 2-segmented (Figs. 4C, D; 6A, B), and legs 4-segmented (Figs. 3E, F; 5E, F). The comparison of the larvae of Pseudolampsis spp. with other described Alticini is difficult because some characters are not clear in the description or/and illustra- tions. A good comparison with Chaetocnema denticulata (Illiger) and C. pulicaria Melsheimer was possible based on the detailed description of Anderson (1938). Comparing the larvae of these two genera, only Pseudolampsis present abdominal segments divided transversally into 2 parts (Figs. 3B, 5C), glandular openings lat- erally on segments 1-8 (Figs 3A, 5B), endocarina short (Figs. 3C, 5G), and cardo glabrous (Figs. 4C, 6A). Other than Pseudolampsis, no larvae are known in the monoplatine genera. Some workers place monoplatines as close to the oedionychine flea beetles possibly based on the swollen hind tarsi and femora both shared (Seeno and Wilcox, 1982). How- ever, Bechyne and Bechyne (1975) separate the oedionychine and disonychine groups from the rest of the Alticini (“Alticidae”) based on apomorphies of the median lobe of the adeagus. Known larva of oedionychine genera Kushelina (Law- son, 1991), Alagoasa (Samuleson 1985, Duckett unpublished) and Asphaera (Duck- ett, unpublished) all lack stemmata and show considerably less sclerotization of the body than larvae of Pseudolampsis. 1997 PSEUDOLAMPSIS 63 ACKNOWLEDGMENTS We are grateful to Cleide Costa of the Museo de Zoologia de Universidade do Sao Paulo (MZSP) for her offer of material of Pseudolampsis darwini and for bringing this problem to our attention, and to Gary Buckingham for this generous gift of P. guttata material, without which this study could have been much poorer. The second author thanks Vilma Savini; MIZA- Museo del Instituto de Zoologia Agricola, Maracay Venezuela, Philip Perkins; MCZ- Museum of Comparative Zoology, Harvard, and Sharon L. Shute; BMNH- The Natural History Museum, London for their help and many kindnessess during visits to collections in their care. She also thanks David G. Furth, Smithsonian Institution, for discussion and comparison with the material in his collection. Terry Seeno and R. Wills Flowers kindly supplied needed bibliography. Com- ments from Wills Flowers and an anonymous reviewer greatly improved the manuscript. Maria Berio and Aurora Lauzardo kindly helped translate the abstract. We thank Frances L. Fawcett for the habitus drawing and Julio Garay for genitalic drawings. This study was partially sup- ported by Conselho Nacional de Desenvolvimento Cientifico e Technoldgico (CNPq) to S.A.C. and by Universidad de Puerto Rico Grants “Fondos Institucionales para Investigacion” (FIPI) numbers 8-80-624, and 8-80-716 to C.N.D. LITERATURE CITED Anderson, W. H. 1938. Description of the larvae of Chaectonema denticulata (Illiger) and Chaectonema pulicaria Melsheimer (Coleoptera, Chrysomelidae). Proc. Entomol. Soc. Washington 40:161-169. Balsbaugh, E. U. 1969. Pseudolampsis (Coleoptera: Chrysomelidae, Alticinae): distribution and synonymy). Coleopt. Bull. 23:16-18. Buckingham, G. R. and M. Buckingham. 1981. A laboratory biology of Pseudolampsis gutata (LeConte) (Coleoptera: Chrysomelidae) on waterfern Azolla caroliniana Willd (Pteri- dophyta: Azollaceae). Coleopt. Bull. 35(2): 18 1-1 88. Chapuis, P. 1875. In Lacordaire, Histoire naturelle des insectes. Genera des Coleopteres. Vol. 11, Eamille des Phytophages, 420 ppl, pis 124-134. Paris Clark, H. 1860. Catalog of Halticide in the collection of the British Museum. Part I, 301pp. London. Plowers, R. W. and D. H. Janzen. 1997. Feeding records of Costa Rican leaf beetles (Cole- optera: Chrysomelidae). Florida. Entomol. (In press). Elowers, R. W. and R. S. Tiffer. 1992. Comportamiento gregario de la vaquita Hypolampsis sp. (Coleoptera: Alticinae) en Guanacaste, Costa Rica. Brenesia 37:135-136. Eurth, D. G. 1988. The jumping apparatus of flea beetles (Alticinae)- the metafemoral spring. In: P. Jolivet, E. Petitpierre and T. H. Hsiao (eds.). Biology of the Chrysomelidae. Dr. W. Junk Publ., The Netherlands: 285-297. Habeck, D. H. 1979. Host plant of Pseudolampsis guttata (LeConte) (Coleoptera: Chryso- melidae). Coleopt. Bull. 33(2): 150. Horn, G. H. 1889. A synopsis of the Halticini of Boreal America. Trans. Am. Ent. Soc. 16: 163-320. Jolivet, P. and T. J. Hawkeswood. 1995. Host plants of the Chrysomelidae of the World. An Essay about the relationships between the leaf-beetles and their food plants. Backhuys Publishers, Leiden, The Netherlands. 281pp. Konstantinov, A. S. 1994. Comparative morphology and some evolutionary trends in flea beetles (Alticinae). In: P. H. Jolivet, M. L. Cox and E. Petitpierre (eds.). Novel aspects of the Biology of the Chrysomelidae, Kluwer Academic Publishers, the Netherlands: 383-391. Lawrence, J. E and A. E Newton, Jr. 1995. Eamilies and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names, pp. 779-1006 In: J. 64 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Pakaluk and S. Slipinski (eds.), Biology, phylogeny and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Warsawa, Museum i Instytut Zoologii PAN. 2:559-1092. Lawson, E A. 1991. Chrysomelidae (Chrysomeloidea) (=Cassididae, Cryptocephalidae, Me- galopodidae, Sagridae, etc.). In: E W. Stehr (ed.). Immature Insects 2 Kendall/Hunt Publishing Company, 975 pp. LeConte, J. L. 1884. Short studies o North American Coleoptera. No. 2. Trans. Am. Ent. Soc. 12:1-32. Leng, C.W. 1920. Catalog of the Coleoptera of America, north of Mexico, Mount Vernon, N.Y., 470 pp. Reid, C. A. M. 1992. The leaf-beetle genus Microdonacia Blackburn (Coleoptera, Chryso- melidae, Galerucinae): revision and systematic placement. Syst. Entomol. 17:359-387. Reid, C. A. M. 1995. A cladistic analysis of subfamilial relationships in Chrysomelidae sensii lato (Chrysomeloidea). pp. 559-631 In: J. Pakaluk and S. Slipmski (eds.). Biology, phylogeny and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Warsawa, Museum i Instytut Zoologii PAN. v 2:559-1092. Samuelson, G. A. 1985. Description of a new species of Alagoasa (Coleoptera, Chrysomelidae) from southern Brazil associated with Lantana (Verbenaceae). Rev. Bras. Entomol. 29: 579-585. Scherer, G. 1962. Bestimmungsschliissel der neotropischen Alticinen Genera (Coleoptera: Chrysomelidae: Alticinae). Entomol. Arb. Mus. G. Erey. 13(2):5 1 1-527. Scherer, G 1964. Eine neue Distigmoptera, die Charles Darwin wahrend seiner Reise auf der “Beagle” fing. Entomol. Arb. Mus. G. Erey. 15:296-301. Seeno, T. N. and J. A. Wilcox. 1982. Leaf Beetle Genera (Coleoptera: Chrysomelidae). En- tomography 1 : 1 -22 1 . Received 15 October 1996; accepted 3 July 1997. J. New York Entomol. Soc. 105( 1— 2):65— 104, 1997 TYPE SPECIMENS OF LEPIDOPTERA IN THE TEPPER COLLECTION AT MICHIGAN STATE UNIVERSITY John H. Wilterding Department of Entomology, Michigan State University, East Lansing, MI 48824 Abstract. — Holdings of Lepidoptera type material in the Tepper collection at Michigan State University are completely evaluated and annotated. Acquired in 1889, the collection contains types of 180 nominate Lepidoptera, predominately noctuids described by H. K. Morrison. A brief curatorial history of the collection as well as handwriting examples for authors of the type material in the collection is presented. The types are completely annotated with bibliographic citation, type locality, condition, and a complete discussion of the status of each type. The following Morrison lectotypes are also designated: Tarache crustaria, Agrotis decolor, Calo- campa germana, Hadena congermana, Agrotis acclivis, Segetia orbica, Tarache patula, and Eurois astricta. Key words: Lepidoptera, Types, Tepper, Cook, Morrison, Lectotypes. For slightly over 100 years Michigan State University (MSU) has been the re- pository of the Tepper Collection of Lepidoptera. The collection of over 12,000 specimens (8,000 species) of butterflies and moths was purchased in 1889 from Fred Tepper of Brooklyn, New York. Included in this collection were a number of type specimens of mostly North American Macrolepidoptera; the majority being species described by Mr. H. K. Morrison. Although biographical information on Tepper is lacking, we know that Tepper was an avid collector, was active in his interactions with specialists and was an early member of a nucleus of individuals who formed the Brooklyn Entomological Society (Franclemont, pers. comm. 1994). He also purchased the collection of H. K. Mor- rison, probably sometime in 1876, before Morrison’s departure to California and the Washington Territory to collect Lepidoptera. The collection contains 180 types (or putative types), the vast majority of these described by H. K. Morrison. In addition, a lesser number of types of Edwards, Grote, Hulst, Smith and Tepper are found in this collection. The majority of the types are noctuids, but sessiids, geometrids, sphingids, notodontids, saturniids, and limacodids are also represented. Over the years, this collection has been an important resource for workers of North American Lepidoptera. It has also been the source of some of their frustrations; this project in part is an attempt to ameliorate the problem by presenting an accurate account of the types in the Tepper collection. The collection was acquired by MSU in 1889, through the efforts of then professor of Entomology A. J. Cook. In the summer of 1888 Cook learned from “a noted professor in one of our Eastern colleges” (probably J. B. Smith of Rutgers) that the Tepper collection was for sale (Anonymous, 1889). Shortly after. Cook and then MSU President Willits approached the State Board of Agriculture with an appeal for the purchase the collection, for $5,000. Senator James MacMillan, upon hearing of the request, wrote Cook stating “I have considered the matter of the Tepper Collec- 66 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) tion, and have decided to authorize you to make the purchase of this collection of insects” (Anonymous, 1889). Cook then traveled to New York to arrange for its shipment and the collection arrived at Michigan State on March 5, 1889. I began this study on the request of Fred Stehr that I “quickly catalog” the Lepidoptera types that we had in the MSU collection to be placed in a separate part of the collection. It soon became clear that there were numerous curatorial mistakes and inconsistencies, many that were quite serious. These errors included the loss of the original combination associated with some specimens, loss of the specific epi- thets, non-type specimens mixed in with the type series, inaccurate dates, misspell- ings, etc. It was also believed that there might be types “hiding” in the general Lepidoptera collection that were not recognized as types. This project was undertak- en because it was almost certain that if a lepidopterist did not soon attend to these errors, these problems would only worsen with time. It was clear that an accurate accounting of the type material in the Tepper collection was long overdue and that the holdings of types in the collection should be accurately communicated to the research community before knowledge of the location of a type, or type specimens, was lost. Moreover, potential changes in the Zoological Code of Nomenclature ne- cessitate that future workers have the best opportunity to study all relevant type material, since the conclusion of the last principle reviser as to the application of names in synonymy will take precedence following publication (Terry Erwin, pers. comm. 1994). EARLY CURATION When the Tepper collection was received at MSU, Professor A. J. Cook made an effort to record all the species (and number of specimens) received in a notebook. The presence of this catalogue was brought to my attention by M. C. Nielsen, and is still in our possession. On the first page of these notes, on MSU Stationery of that period, is written “Catalogue of the MacMillan Collection of Lepidoptera made in 1889 from the collection as received from Mr. Tepper.” These notes proved in- valuable because they indicate which species in the collection were represented by types. It appears that based on this typed inventory of the Tepper collection, each type specimen was labeled, perhaps by Cook, with a unique yellow, rhomboidal label (Fig. 1). Apparently, no effort was made to add information to this material other than the addition of the yellow rhomboidal type label. Therefore, if a type was Fig. 1. Example of the type label and handwriting of A. J. Cook, from the type of Litha- codia penita Morrison. Fig. 2. Example of Frederick Tepper type label from the type of Drynobia tortuosa Tepper. Tepper locality label from type of Oncocnemis riparia Morrison. Handwriting and typical label shape of Morrison from the type of Mamestra teligera Fig. 3. Fig. 4. Morrison. Fig. 5. Fig. 6. Fig. 7. Fig. 8. Type label of Agrotis personata Morrison, atypical of Morrison’s label design. Type label and handwriting example from the type of Heliophila patricia Grote. Type label and handwriting from the type of Nemoria tepperaria Hulst. Type label of Mamestra dodgei Morrison in an unknown hand, perhaps Dodge. 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 67 68 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) without a label indicating either the author or the specific epithet, it was to remain this way. While this situation may seem chaotic to us today, in the late 19th century, type concepts were in a state of flux, and curation of types, and placement data on specimens was far from standardized. Therefore, unless the authors did so them- selves, the specimens often did not have the original name, or the word “type” associated with the specimen (Franclemont, pers. comm. 1994). In the case of the Tepper collection, this has resulted over the years in the inaccurate curation, and even worse, the “loss” of type material. In more than a few cases the recovery of ‘Tost” types has been accomplished by a careful search of the general collection. We know that initially the collection was housed separately from the main col- lection in “MacMillan Cabinets” (Anonymous, 1889). Some time later, this material was interpolated into the general collection as the collection began to expand. The types remained scattered throughout the general collection until the 1950’s when “all” the types were pulled from the collection by Roland Fischer and housed sep- arately. But only those specimens with the rhomboidal “type” label were removed; any material not so labeled, or with a hidden “type” label, was apparently over- looked. In an effort to recapture any potential “lost” types, I searched through the entire collection looking for types (mostly Morrison) which may have been over- looked in the past. BACKGROUND AND TREATMENT OE THE TYPES There are a number of features of the type collection worthy of careful explanation that were not fully appreciated by earlier workers of the material. In nearly all cases at least one specimen of an original type series bears the yellow rhomboidal “type” label (Fig. 1) that was erroneously interpreted by many as a Tepper “type” label (Todd, 1982). This label, however, is probably in Cook’s, not Tepper’s hand; ex- amples of Tepper’s handwriting can be seen in Figs. 2-3. Through careful study of the entire collection, it was discovered that this rhomboidal label was also associated with other non-lepidopterous type material described from the collection in the 19th century, and also matches the hand of the individual who wrote “type” in the orig- inal accession notes. It seems likely that this label was added after the collection was received at MSU, and was likely done so by Cook or other worker and appears to have been a general policy in the treatment of all types in the collection near the turn of the century. Except in a few instances, regardless of the number of specimens in the type series, only one specimen in a type series is labeled “type” with this rhomboidal label. This has resulted in the inclusion or exclusion of additional specimens from the type series and has necessitated referring to the original descriptions in order to clearly define the limits of the original type series. Morrison’s types comprise the bulk of the Lepidoptera type collection. This is fortunate because Morrison usually adequately labeled the specimens with the spe- cific name and “Morr.” He usually employed a distinctively shaped label (Fig. 4), that he rarely deviated from (Fig. 5) and which frequently included the location and date of capture. In all but a few cases, only one specimen in an otherwise syntypic series has this detailed label. Other specimens in a type series often lack data labels of any kind, or more frequently, only the state or territory of capture. Based on 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 69 Morrison’s type labeling protocol, there is some evidence that suggests that Morrison had an operational holotype concept at this time, in part due to his practice of selecting only one specimen for this detailed label. His specimens reflect this well, for where there is more than one in a type series, only a single specimen is labeled with the word “type” on his peculiar label. Other authors have described species from material in the Tepper collection, and more often these were originally described from Morrison’s collection before it was purchased by Tepper. Smith described a number of noctuids from the Tepper collec- tion, some of which Todd (1982) recognized as valid types in his work on the Smith type material. Unfortunately, he seems to have missed a number of types and his- torical specimens with notes written by Smith which were still “hidden” in the general collection. I have not endeavored to recapture type material that Todd over- looked since lectotypes and type identities were fixed for all Smith names in Todd’s work. Todd (1982) designated a number of Tepper types as lectotypes, but in only one case was a lectotype label affixed to the specimen; lectotype labels have since been added to the proper specimens based upon Todd’s publication. In addition, types of Grote (Fig. 6), Tepper (Figs. 2,3), and Hulst (Fig. 7), can be found in the type collection. In addition, there are several data labels whose authorship I have been unable to trace (Fig. 8). There were a number of sources used to assist in the recovery of additional type material and to confirm the present location of many types. Smith (1893), in his Catalog of Lepidoptera, discussed the location of the type, and whether he had examined it or not. Taxa which he explicitly mentioned having seen in the Tepper collection, were cross-referenced with the type, and general collections. The general collection has also been checked against all (or most) of the Morrison names re- sulting in the discovery of 28 unlabeled types (some putative types). Many of these types were previously believed lost. Poole’s (1989) catalog of the Noctuidae was also used for clues as to the location of type material, recognizing Poole’s assertion that he often was not certain of the location of types, but was making educated guesses. I have attempted to be as accurate as I can when making decisions as to the number of types in the original series and the validity of “type” material. It was not uncommon during this period for a new species to be described from a number of specimens from private collections. Usually these specimens were returned to the private collector; and therefore types for a single name may presently be found in a number of collections. Therefore, for example, it is not uncommon to expect material to be both at MSU, and the AMNH. These non-MSU specimens were not tracked at other institutions, but published lists and access to Internet data bases were used to check type holdings at other museums. On several occasions lectotypes are designated, and the rationale for doing so is stated in the species accounts; I have endeavored to do so only when there are no closely related species and when it is reasonably certain that there is not additional type material at other institutions. In addition, lectotypes are not designated in tax- onomically difficult genera or species groups. At no time in the work do I make nomenclatorial decisions, or take such action as removing specimens from the type series when clear evidence was not found to support such action. My primary goal 70 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) in this work is to communicate exactly what is in the collection so that future workers might be better informed when studying this collection. The types are listed alphabetically by species name within each family. For each type(s) the original combination, citation, and a quote from the description as to the type locality and, when stated, the source (collector) of the specimen is given. Also included is a list of specimens noting the sex and the data from each label listed in order (separated by “”) from highest to lowest label on the pin with notes or ques- tionable letters in brackets []. After each label the author’s handwriting is indicated in brackets []: (M = Morrison, T = Tepper, H = Hulst, C = Cook (yellow rhom- boidal label), G = Grote, ty = type print, S = Smith). Horn and Kahle (1935-1937) was used to confirm the handwriting identity, however, this was not possible in the case of Tepper, Cook, or Morrison. The condition of the specimen is given following label information. For each species I give relevant remarks pertaining to the material, my conclusion as to the validity of the type material, the present combination and the Hodges (1983) number (RWH). In some instances there was non-type material mixed with the type(s); these specimens have been removed, and no indication of doing so is made when such action was not ambiguous. I have referred to, whenever possible, publications by the last principle reviser to cross check conclusions that they may have reached regarding the location of type material. The material is available for study by qualified specialists, and we encourage the use of this valuable type collection in future revisionary projects. Workers should be aware that most of the Tepper material is still extant, in the general collection. I have gone through much of the collection trying to recover the Morrison types, however, the larger task of cross-checking all the Grote names with their original descriptions, and the possibility of overlooked Morrison and Hulst types remains a possibility. It is likely that not all of the types have been retrieved from the general collection. Suggestions as to other types that we should search for are welcome. Sesiidae Aegeria albicornis Hy. Edwards, 1881b:201. Type locality. “1 male. Nevada. (Morrison). Type. Coll. F, Tepper.” Male, “Nevada” [T]; “Type”[C]; “Female” [symbol] [In good condition] Remarks: The holotype (Englehardt, 1946). Synanthedon (RWH 2570). Aegeria brunneipennis Hy. Edwards, 1881b: 191. Type locality. “1 female. Georgia. (Morrison). Type. Coll. F. Tepper.” Female, “Ga”; “F. T.”[T]; “Type”; “Female” [symbol] [Type in poor condi- tion; the abdomen and legs missing, the left antenna is broken near middle] Remarks: The holotype. Englehardt (1946) gives no indication as to the location of the type. Eichlin and Duckworth (1988) give MSU as the respiratory. A synonym of Synanthedon rileyana (Hy. Edw.) (RWH 2552). Sciapteron graefi Hy. Edwards, 188 lb: 183. Type locality. “1 male. 2 females. Nevada (Morrison.)” “Type. Coll. E. L. Graef Female, “Nevada” [T]; “F. T.”[T] “Type”[C]; “Female” [In good condition] Female, “Nevada” [T]; “F. T.”[T]; “Type”[C]; “Female” [In good condition] 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 71 Remarks: Syntypes. Englehardt (1946) and Eichlin and Duckworth (1988) both state that the LI.S.N.M. has the type. However, it is likely that the two females at MSU are also types and were obtained by Tepper through his purchase of the Mor- rison collection. A synonym of Synanthedon exitosa (Say) (RWH 2583). Pyrrhotaenia helianthi Hy. Edwards, 18815:203. Type locality. “1 Male, 1 female. Virginia City, Nevada. (H. E.) On Helianthus, sp. Types. Coll. Hy. Edwards.” Female, “Nevada” [T]; “E T.”[T]; “Female” [Specimen in very good condi- tion] Remarks: Englehardt (1946) states that the type is in the AMNH. Eichlin and Duckworth (1988) state the type is at MSU. This is probably the holotype. A syn- onym of Synanthedon polygoni (Hy. Edw. 1881) (RWH 2581). Aegeria inusitata Hy. Edwards, 18815:201—202. Type locality. “White Mts. N. H. (Morrison.) Andover, Mass. (F. G. Sanborn.)” “Types. Coll. F. Tepper.” Female, “W. Mts. N. H.”[M]; “Type”[C]; “female” [symbol] [In very good condition] Male, “W. Mts. N. H.”[M] “86”[?] “male” [symbol] [In very good condition] Remarks: Undoubtedly part of the original syntype series. Engelhard! (1946) makes no reference to the location of the type(s) of this name. Eichlin and Duck- worth (1988) make reference only to the Andover specimen which is in the AMNH. They make no reference to the location of the NH specimen(s). A synonym of Synanthedon pictipes (G.&R.) (RWH 2550). Carmenta minuta Hy. Edwards, 18815:185. Type locality. “1 male. Georgia. (Morrison.) Type. Coll. F. Tepper.” Male, “Georgia” [T]; “Type”[C]; ''minuta Hy Ed”[M]; “female.” [symbol] [In poor condition, the abdomen and right forewing missing] Remarks: The holotype (Eichlin and Duckworth 1988). A synonym of Osminia ruficornis (Hy. Edw.) (RWH 2545). Albuna montana Hy. Edwards, 18815:188. Type locality. “21 examples.” “White Mountains, N. H. (Morrison. Grote.) Ne- vada, Colorado. (Morrison.) Anticosti Isld. (Couper.) Ser. Nevada, Cal. (H. E.). Colls. Tepper, Graef, Neumoegen, W. Grey, Dr. Bailey, Hy. Edwards, etc., etc.” Female, “W. Mts. N. H.”[T]; “F. T.”[T]; “Type”[C]; “female” [symbol] [In good condition, the meso legs missing] Male, “W. Mts. N. H.”[T]; “F. T.”[T]; “male” [symbol] [In good condition, the right meso and meta leg missing] Male, “W. Mts. N. H.”[T]; “F. T.”[T]; “male” [symbol] [In good condition] Male, “Anticosti” [T]; “F. T.”[T]; “male” [symbol] [In poor condition, antennae missing, meso legs missing, considerable development of verdigris] Female, “W. Mts. N. H.”[T]; “F. T.”[T]; “female” [symbol] [Damage by der- mestids to the abdomen, and antennae broken near base] Male, “Mt. Hood”[T]; “male” [symbol] [In fair condition, the left meso leg missing] Remarks: Syntypes. Englehardt (1946 p. 165) states that “the type (a single fe- 72 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) male) is in the American Museum.” Eichlin and Duckworth (1988) state the type from Colorado is in the AMNH. A synonym of Albuna pyramidalis (Wlk.) (RWH 2533). Aegeria opalescens Hy. Edwards, 1881b: 199. Type locality. “3 males. Virginia City, Nevada. (H.E.).” “1 female. Colorado. (Morrison.).” Male, “Nevada” [T]; “F. T.”[T] [In fair condition, the pro and meso legs miss- ing] Male, “Nevada” [T]; “E T.”[T] [In good condition] Remarks: Syntypes. The collection should have the Morrison female from Colo- rado, but I have been unable to locate it. Eichlin and Duckworth (1988) state that the type from Nevada is in the USNM; these two males are likely not part of the original syntypic series. A synonym of Synanthedon exitosa (Say) (RWH 2583). Pyrrhotaenia orthocarpi Hy. Edwards, 1881b:204. Type locality. “3 males. 1 female. On Orthocarpus luteus. Nutt. Washoe Lake, Nevada, (H. E.). “Type Coll. Hy. Edwards.” Male, “Nevada” [T]; “male” [symbol] [In good condition, left meta and right meso leg missing] Male, “Nevada” [T]; “male” [symbol] [In very good condition] Male, “Nevada” [T]; “male” [symbol] [In good condition, antennae are missing] Remarks: The status of these specimens is not known. No mention was made by Englehardt (1946) as to location of the type of this species. Eichlin and Duckworth (1988) state that the type is in the AMNH. A synonym of Synanthedon polygoni (Hy. Edw.) (RWH 2581). Aegeria proxima Hy. Edwards, 188 lb: 201. Type locality. “White Mts. N. H. (Morrison.) Type. Coll. F. Tepper.” Male, “W. Mts N. H.”[T]; “E T.”[T]; “Type”[C] [In good condition, left an- tenna missing] Remarks: The holotype (Engelhardt 1946) and (Eichlin and Duckworth, 1988). Synanthedon proxima (Hy. Edw.) (RWH 2572). Aegeria seneciodes Hy. Edwards, 1881b:198-199. Type locality. “1 male. California. On Senecio douglassii. D. C. (H. E.) 1 female. Nevada. (H. K. Morrison.) Types. Coll. Hy. Edwards. E Tepper.” Male, “Nevada”[T]; “E T.”[T]; “Female” [symbol] [Left pro and meso and right meso leg missing; otherwise in fair condition] Remarks: Engelhardt (1946) states that the male type of Aegeria seneciodes is in AMNH. Eichlin and Duckworth (1988) stated the type from California is in the AMNH. This is also a type. A synonym of Synanthedon mellinipennis (Bdv.) (RWH 2580). Pyrrhotaenia tepperi Hy. Edwards, 1881b:203. Type locality. “1 male. Georgia, (Morrison.)” “Type. Coll. F. Tepper . . .” Female, “Georgia” [T]; “Type”[C]; “female” [symbol] [In poor condition, the antennae are glued to the frons, the legs missing, and portions of the wings torn] 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 73 Remarks: Engelhard! (1946) and Eichlin and Duckworth (1988) state that the (holo)type is in MSU. A synonym of Synanthedon acerni (Clem.) (RWH 2554). Aegeria verecunda Hy. Edwards, 188 lb: 190. Type locality. “1 male. 2 females. Colorado. (Morrison.)” “Type. Coll. Hy. Ed- wards. E Tepper. Female, “Col”[T]; “F. T.”[T]; “Type”[C]; “female” [symbol] [In good condi- tion] Remarks: The holotype (Eichlin and Duckworth 1988). Carmenta (RWH 2615). Urodidae Penthetria parvula Henry Edwards, 188 la: 80. Type locality. “1 male, Indian River, Florida, 1 female, Georgia.” “Types Coll. Hy. Edwards. E Tepper.” Male, “Georgia” [T]; “Type”[T] [Type in very good condition] Remarks: Syntype. The sex of the specimen from Georgia in the original descrip- tion was probably misdetermined. Urodus (RWH 2415). Geometridae Acidalia cacuminaria Morrison, 1874E195. Type locality. “Massachusetts, about the first of July, Collection of H.K. Morri- son.” Female, ^^cacuminaria Morr. Type, Mt. [H]om Ms. July 6, 79” [M] [In fair condition, abdomen glued, upside down (Coveil 1970). Remarks: The holotype (Covell, 1970). Scapula (RWH 7157). Endropia decoloraria Hulst, 1886:207. Type locality. “2 males Coll. Graef and Tepper. No locality with either specimen.” Male “Type”[H] [In poor condition, the abdomen missing, only the right pro leg present] Remarks: Syntype. This is a valid type specimen; the other male type is in the AMNH (Rindge, 1955). Cepphis (RWH 6834). Drepanodes ejfascinaria Hulst, 1886:204-205. Type locality. “2 females S. C. and Fla: coll. Neumoegen and Tepper. 42 mm.” Female, “S. Car.”[?], “Type”[H] [In fair condition, the abdomen glued to the thorax] Remarks: Syntype. The female syntype from South Carolina is in the AMNH (Rindge, 1955). A synonym of Eusarca fundaria (Gn.) (RWH 6933). Boarmia floridaria Hulst, 1886:215. Type locality. “1 male, 1 female, Fla.” Female, “Type”[H]; “Florida” [T] [Abdomen and antennae missing] Remarks: Syntype. There is no indication in the original type series as to the source of the material. The AMNH has the male syntype from Florida (Rindge, 1955). A synonym of Idaea violacearia (Wlk.) (RWH 7120). Boarmia furfuraria Hulst, 1888:214. 74 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Type locality. “3 males. Col.” “Expands 42mm.” Male, “Col”[T]; “Type”[H] [In good condition, the meta legs and the left antenna missing] Remarks: Paralectotype. The lectotype of this species from Colorado is in the AMNH (Rindge, 1958). Glena (RWH 6446). Chlorosea graefiaria Hulst 1886:123. Type locality. “2 females, Nevada, Coll. Graef, Tepper.” Female, “Nevada” [T]; “Type”[H] [The well marked specimen missing both meta and the right meso thoracic legs] Remarks: Syntype. The Graef specimen is in the USNM (Ferguson, 1985). Che- teoscelis (RWH 7068). Endropia helveolaria Hulst, 1881:33-34. Type locality. “One specimen, also one in collection of Mr. Fred Tepper, Colorado. Male, “Col”[T] [In good condition] Remarks: This syntype was found in the general collection. A male syntype from Colorado is also in the AMNH (Rindge, 1955). Lychnosea (RWH 6857). Endropia hilumaria Hulst, 1886:206-207. Type locality. “2 males Colorado. Coll. Graef, Tepper.” Male, “Col”[T]; “Type”[H] [In very good condition] Remarks: Syntype. There is also a syntype of this species in the AMNH (Rindge, 1955). Caripeta (RWH 6871). Endropia manubiaria Hulst, 1886:207—208. Type locality. “2 males, 1 female Colorado. Coll. Tepper, Hulst.” Male, “Col”[T], “Type”[H] [In poor condition, the right wings, both antennae, and the right meso and meta legs destroyed] Remarks: Syntype. Euchlaena (RWH 6736). Tetrads mellitularia Hulst, 1886:202-203. Type locality. “Five males, 3 females. Ariz., Nev., Cal; Coll. Hy. Edwards, Tepper, Hulst.” Male, “Nevada” [T] [In very good condition] Remarks: Paralectotype. The lectotype is in the AMNH (Rindge, 1955). A syn- onym of Pharene placeraria (Gn.) (RWH 6949). Endropia occantaria Hulst, 1886:207. Type locality. “Two females Nev.: Coll. Graef, Tepper.” Female, “Nevada” [T] [In very poor condition, extensive dermestid damage to the abdomen and thorax, head missing] Remarks: The holotype. Rindge (1955) reported from AMNH “a female labeled “type” from [the] Washington Territory. This is probably spurious, because the type locality is said to be Nevada.” A synonym of Euchlaena mollisaria (Hulst) (RWH 6730). Hybernia olivacearia Morrison, 1874f[1875]:200. Type locality. “Massachusetts. From April 20th to May 5th.” Male, ''olivacearia Morr. Type. Cambridge. M[S] 4. 30 155” [M] [In very good condition] 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 75 Male, “5.20” [M]; “Cambridge” [M] [In excellent condition] Female, “4/25” [M]; “Cambridge” [M] [In very good condition] Remarks: Paralectotypes. Rindge (1955) designated a male in the MCZ as the lectotype. A synonym of Phigalia strigataria (Minot) (RWH 6660). Tornos robiginosus Morrison, 1874g[1875]:218. Type locality. “Waco, Texas.” Female, “Bastrop Texas” [M] [Right antenna missing, otherwise in good con- dition] Remarks: Paralectotype. This specimen was found in the general collection. I have dissected this specimen and it is scolopacinarius. The lectotype is in the USNM (Rindge, 1954). Tornos scolopacinarius (Gn.) (RWH 6486). Nemoria tepperaria Hulst 1886:122. Type locality. “1 male, 1 female, Ga. N. Car. Coll Edwards, Tepper.” Male, “Ga”[H]; “Type”[H] [The type is in poor condition. The antennae are broken near base, the right pro and meso and left pro legs are also missing] Remarks: Syntype. This male was part of the original type series. Ferguson (1985) stated that the (syn)type from N. Car. is in the USNM. Chloropteryx (RWH 7075). Tephrosia texanaria Hulst, 1888:216. Type locality. “Eight males, six females.” Male, “Tex.”[ty]; “Type”[H] [Abdomen and antennae missing] Male, “Tex.”[ty] [The well marked specimen is in good condition, only the right prothoracic leg is missing] Remarks: Paralectotype. The original description gave no information on the lo- cation of the types, but the male marked “Type” is in Hulst’s hand and is typical (red border) for his type labels. Given the large number in the original series, it is likely that the other males marked with the same small “Tex” and the other without information are also part of the type series. The lectotype of this species was des- ignated by Rindge (1973). Glenoides (RWH 6443). Boarmia wrightiaria Hulst, 1888:215. Type locality. “4 females.” “Taken at San Bernardino, Calif.” Male, “So. Cal.”[ty]; “Type”[H] [In poor condition, only the left meta leg present, the right antenna missing and the abdomen destroyed by dermestids. The genitalia appear to be intact] Remarks: Paralectotypes. The lectotype of this species is in the AMNH (Rindge, 1955, 1970). Hulstina (RWH 6547). Limacodidae Parasa fraterna Grote, 1881c:4-6. Type locality. “N.Y., Mass., Mr. Roland Thaxter; also in Mr. Tepper’s collection.” Female, “Type”[T] [Very good condition] Male, “ex L. June 17, 74.” [T]; “Type” [T] [Very good condition] Remarks: Syntypes. A synonym of Parasa chloris (H.-S.) (RWH 4698). Saturniidae Attacus cinctus Tepper, 1883:65-67, 1 plate. 76 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Type locality. “Eight specimens served as types. Southern Arizona.” Female, “Arizona” [T]; “Original Type”[T] [In fair condition, right antenna missing, left meso and meta legs missing] Female, “Arizona” [T]; “Original Type”[T] [In poor condition, dermestid dam- age to the abdomen] Remarks: Paralectotypes. The lectotype was designated by Ferguson (1972). Rothschildia cincta (RWH 7760). Sphingidae Sphinx albescens Tepper, 1881:1-2. Type locality. “One male from Colorado, taken by Mr. H. K. Morrison, in my coll.” Female, “Col”[T]; “Type”[T] [In poor condition, the abdomen is partially de- stroyed by dermestids, and reglued to the specimen] Remarks: The holotype. This type was found in the general collection. A synonym of Sphinx vashti (Stkr.) (RWH 7803). Notodontidae Drynobia tortuosa Tepper, 1881:2, plate 2 figure 2. Type locality. “One male from Colorado collected by Mr. H. K. Morrison. Type in my collection.” Male, “Col”[T]; “Type”[T] [In very good condition] Remarks: This holotype was found in the general collection. Hyperaeschra (RWH 7918). Noctuidae Charadra decora Morrison, 1875:55. Type locality. “California.” Female, “Calif” [T] [In poor condition, left antenna, left and right meta legs missing, dermestid damage to abdomen] Remarks: The holotype. This specimen was found, unlabeled, in the general col- lection. Lichnoptera (RWH 9187). Agrotis acclivis Morrison, 1875L93-94. Type locality. “New York from the Tepper collection.” Male, “Long Island 191 ”[M]; “Type”[C] [In fair condition; some dermestid damage, right antenna missing] Male, “1706”; “Type”[C] [In fair condition; both meso legs missing, the right antenna and the valves partly destroyed] Remarks: The male, from Long Island, is selected, labeled, and designated as the lectotype. Loxagrotis (RWH 10870); Richia Poole (1989). Hadena adnixa Grote, 1880a:243. Type locality. “Nevada; Tepper. Expanse 38 mil.” Male, “Nevada” [T]; “Type”[T] [In very good condition, the right antenna missing] 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 77 Remarks: This specimen was found in the general collection with a ‘type’ label hidden under the first label. Poole (1989) stated the type was at the BM, but Smith (1893) indicated the type(s) was in the Tepper collection. In the absence of additional Tepper type material, this is likely the holotype. Aseptis (RWH 9533). Hadena albina Grote, 1874b: 157. Type locality. “California (Mr. Behrens, No. 78, Sauzalito, May 15). Two speci- mens.” Female, “Calif” [T]; “Type”[C] [In good condition; the left meso and meta and the right pro and meta legs missing] Remarks: Smith (1893 p. 138) states that “the ‘type’ is in the British Museum; but I am not sure it is a fair representative of the species Mr. Grote meant to describe. It seems to be really a form of castanea, as Mr. Grote suggested; but the specimen in the Tepper collection also marked type by Mr. Grote, is more nearly allied to arctica, and is a good species as shown in my Revision.” The remark by Smith would suggest that the specimen in MSU was labeled type by Grote, but the presence of the rhomboidal label brings this conclusion into question since this is a Cook label. This makes it uncertain if indeed this specimen was seen by Grote at the time of description. Poole (1989) stated that the type is in the BM, however, in Cook’s accession notes, “type” is written next to the name in his hand. The type status of this specimen remains uncertain, but is likely not a type. A synonym of Apamea castanea (Grote) (RWH 9347). Hypena albopunctata Tepper, 1881:2, plate 1, figure 5. Type locality. “One male from Wash. Ter., collected by Mr. H. K. Morrison, in my collection.” Male, “W.T.”[M]; “Type”[T] [Poor condition, legs and antennae missing] Remarks: The holotype. A synonym of Hypena humuli Harris (RWH 8461). Orthosia americana Morrison, 1875c [1876]:434. Type locality. “Mr. W. V. Andrews of Brooklyn has been so kind as to send me from his collection a specimen taken in N.J.” Male, “N.J.”[?]; “Type”[C] [A well marked specimen in fair to poor condition, dermestid damage to thorax and abdomen, left antenna missing] Remarks: The holotype. A synonym of Agrochola lota (Clerck) (RWH 9956). Acronycta aspera Morrison, 1874b[ 1875]: 132. Type locality. “Adirondack Mts., N.Y.” Female, “N.Y.”[T]; ''aspera Morr Type. [A]dr Mts. N. Y. 124” [M] [Type in very good condition] Remarks: The holotype. A synonym of Andropolia contacta (Wlk.) (RWH 9564). Mamestra assimilis Morrison, 1874c: 113. Type locality. “Massachusetts. Collection of H. K. Morrison.” Male, “Tepper” [ty]; “Type”[C] [Specimen in fair condition, right antenna and meso and meta legs missing, valves partly destroyed] Remarks: The holotype (McCabe, 1980). Melanchra (RWH 10295). Enrols astricta Morrison, 1874b[ 1875]: 135. 78 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Type locality. “New Hampshire.” Male, “W. M.”[M] [In good condition] Female, “W. M.”[M] [In fair condition, left antenna missing, right wings broken off] Remarks: These specimens from the White Mountains of NH were found in the general collection. They are likely the types of this species. The male, is hereby selected, labeled and designated as the lectotype of E. astricta. Poole (1989) states that the type is in MSU (RWH 10930). Metahadena atrifasciata Morrison, 1 875c[l 876] :43 1-432. Type locality. “Orono, Maine. (Prof. C. H. Femald).” Female, “Maine” [M]; “121”; “Tepper”[ty]; “Type”[C] [Condition of speci- men is quite poor, considerable damage from verdigris] Male, “Maine” [M?] [Very good condition] Male, “Maine” [T] [Very good condition] Remarks: Syntypes. The two males were found in the general collection. It is uncertain whether these specimens were part of the original series. Smith (1893) states types are in both Tepper collection and BMNH. A synonym of Oncocnemis piffardi (Wlk.) (RWH 10123). Orthosia belangeri Morrison, 1874b[ 1875]: 149. Type locality. “Quebec, Canada.” Male, “Canada” [T]; “Type”[C] [Type in good condition, right antenna, pro and meso legs missing] Remarks: The holotype. Type material for this species was not found at the BMNH (Mikkola, pers. comm. 1992). A synonym of Apamea inficita (Wlk.) (RWH 9369). Agrotis binominalis Smith, 1888a:451. Type locality. “California, Washington Territory.” Male, “W.T.”[M]; “Type”[T] [In poor condition, verdigris damage to the ab- domen, both pro and right meso leg missing] Male, “W.T.”[M]; “Type”[C] [In poor condition] Remarks: Paralectotypes. Todd (1982) found three specimens, two males from WT and a female from California curated as types of binominalis. He chose not to recognize the MSU specimens as types and went ahead and labeled a specimen in the AMNH as the lectotype. What he probably did not realize was that the type of exsertistigma, a female, was curated under a more recent combination, Rhyncagrotis [sic] binominalis. This female, discussed in Todd (1982) under binominalis, is prob- ably the type of exsertistigma. Like many types of the time, no name is affixed to the type. The males above were part of the original type series of binominalis, and are therefore paralectotypes of this name. A synonym of Rhychagrotis exsertistigma (Morr.) (RWH 11047); misspelled as exertistigma in Hodges (1983). Eucalyptera bipuncta Morrison, 1875f:104. Type locality. “Massachusetts. Taken at Belmont, Aug. 17, 1874.” Male, '"bipuncta Morr Type. Cam. Ms. Aug 7 275 ”[M]; “Type”[C] [Type slightly worn, the abdomen and right antenna missing] Remarks: The holotype. The type locality “Belmont” in the original description 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 79 vs. “Camb” on the specimen are not in agreement, as is the date which was likely a transcriptional error. Belmont, however, is very near Cambridge and is probably referable to this specimen, the date discrepancy is likely a transcriptional error. A synonym of Gabara subnivosella bipuncta (RWH 8522a). Agrotis bocha Morrison, 1874b[1875]: 163. Type locality. “Nebraska, Colorado.” Female, “Colorado 202” [M]; “Tepper”[ty] [Right forewing damaged; the left forewing missing and only the left pro leg present] Remarks: This holotype was found in the general collection, unlabeled. M.C. Niel- sen brought this specimen to my attention as a suspected type. Hardwick (1970:156) stated that the “monotype of bocha should be in the Tepper Collection at MSU, but a thorough search has failed to uncover the specimen and it must be presumed lost.” The Colorado specimen was not found in the general collection. Crassivesica (RWH 10913). Luceria burgessi Morrison, 1874c: 109. Type locality. “Tuckemuck Island, near Nantucket. Four specimens taken by Mr. Bigelow, and now in the collection of Edward Burgess and H. K. Morrison.” Male, “Type”[CJ; “Nantucket 43” [M] [Excellent condition] 2 Females, no data [Excellent condition] Remarks: Syntypes. Although Poole (1989) indicated that the types are in the BMNH, Kauri Mikkola (pers. comm. 1992) was unable to find it there; his feeling is that these three specimens comprise what remain of the original type series. Al- though uncommon, sometimes Morrison’s additional specimens do not bear any other data when taken at the same locality. The fourth specimen of the type series must be presumed lost. In a forthcoming publication, Mikkola and Lafontaine will des- ignate the male listed above as the lectotype (Mikkola, pers. comm. 1995). Crymode s (RWH 9378). Heliothis californicus Grote, 1873c: 149. Type locality. “California (Mr. Hy. Edwards, No. 9). Four specimens examined.” Male, ''californicus Grt, Type. California 299” [M] [Left meta leg absent, left antenna broken at base, abdomen whole but excavated by dermestids, genitalia destroyed] Remarks: Paralectotype. A total of six specimens were curated as types for suetus Grote, one of which was the male type of californicus. The Tepper collection ac- cession notes indicate types of californicus were received with the Tepper collection. Smith (1893) state that one type was in Tepper’s collection, another in the BMNH. Hardwick (1958, 1996) designated the specimen in the BMNH as the lectotype. Schinia sueta californica (RWH 11088a). Thalpochares carmelita Morrison, 1875c [1876]:434-435. Type locality. “Dallas, Texas (Mr. Boll).” Male, “Dallas Tex. Boll”[ty]; “Type”[C] [Specimen in excellent condition, verdigris building around the thorax of the specimen] Remarks: Syntype. The original extent of the type series is not known. Poole (1989) states that the type is in the MCZ, but this specimen is also likely a type. A synonym of Eumicremma minima (Gn.) (RWH 9076). 80 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Hadena castanea Grote, 1874b[ 1875]: 156. Type locality. “California (Mr. Behrens, three specimens with the number 10 and 20).“ Male, “Calif.” [T]; “Type“[C] [In very good condition] Remarks: The status of this type is not known. A total of 6 specimens were curated as types for this name, 5 poor specimens were Tepper’s from the Washington Ter- ritories. The species was described from California, and the male so marked is prob- ably the only potentially valid type, however, the absence of the red number labels indicated by Grote in the original description places this conclusion into question. A total of 8 specimens were noted in the Tepper collection accession notes, and “type” is written next to the name in Cook’s hand. Apamea (RWH 9349). Hadena characta Grote, 1880a:243. Type locality. “Nevada; Mr. Tepper.” Female, “Nevada” [T] [In very good condition] Remarks: The status of this type is not known. This specimen was found in the general collection without a type label. Smith (1893) stated that the type was in the BM and that “another typical specimen” was in the Tepper collection. I have not been able to confirm the presence of this type in the BM. Aseptis (RWH 9543). Agrotis comosa Morrison, 1876:238. Type locality. “Colorado T. L. Mead.” Male, “Type”[C]; “Colorado Aug 7 1872 193” [M] [In poor condition; the abdomen and legs and left antenna missing] Remarks: The holotype (LaFontaine, 1987). Euxoa (RWH 10780). Agrotis confusa Smith, 1888a:452. Type locality. “Washington Territory.” Female, “W. T.”[M]; “Type”[C] [In very good condition] Female, “W. T.”[M]; “Type”[C]; “1599”[?] [Extensive damage by verdigris on thorax] Remarks: Paralectotypes. Todd (1982) designated a specimen at the LFSNM as the lectotype. A synonym of Rhynchagrotis insularis (Grt.) (RWH 11048). Hadena congermana Morrison, 1874d:106. Type locality. “A rare species. Hab. New York, and one specimen taken at Bev- erly, Mass., June 24, 1867, by Mr. Edward Burgess ...” Male, “Beverly Ms. June 24, 1867 25”[M]; “Type”[C] [In excellent condition, left antenna missing] Female, “N.Y. State” [M] [In excellent condition] Male, “N.Y. State” [M] [In poor condition, extensive dermestid damage to the thorax, head missing] Remarks: The data of the male from MA matches perfectly with the original description; it is selected and hereby designated as the lectotype. Sideridis (RWH 10266). Taeniocampa confluens Morrison, 1874b[ 1875]: 159— 160. Type locality. “St. Louis, Mo. Prof. C. V. Riley.” Female, ''confluens, Morr Type, Cen. Missouri 413”[M] [In poor condition, antennae missing, abdomen and thorax severely excavated by dermestids.] 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 81 Remarks: The holotype. A synonym of Orthosia hibisci (Gn.) (RWH 10495). Homophoberia cristata Morrison, 1875b: 125. Type locality. “Hoboken, N.J. One specimen kindly presented to us by Mr. Her- man Sachs.” Female, “N.J.”[?] [In very good condition, the antennae missing] Remarks: This specimen was found unlabeled in the general collection and is probably the holotype of cristata (RWH 9056). Tarache crustaria Morrison, 1875d:70-71. Type locality. “Nebraska. Described from material given us by Mr. G. M. Dodge.” Male, “Type”[C]; “Type Nebraska 258” [M] [In good condition] Male, “Nebr”[?]; “Type”[C] [In poor condition; the specimen damaged by dermestids, the abdomen and wings glued onto the specimen] Remarks: The male in good condition marked type by Morrison is selected, la- beled and now designated as the lectotype. Pseudacontia (RWH 10172). Mamestra curta Morrison, 1875L96. Type locality. “Colorado, July 20 and 22.” Male, “Colorado” [T]; '"curta, Morr, Type. Colorado. 7. 22 220” [M] [A well marked specimen in poor condition; dermestids appear to have excavated the thorax and abdomen but the genitalia appear to be intact. The legs are absent and the abdomen has been glued at the base of the thorax.] Female, "curta Morr Type. Colo July 20 220” [M] [In very good condition] Remarks: The holotype. A subspecies of Lasiestra impingens (RWH 10339a). Agrotis decolor Morrison, 1874b[ 1875]: 162. Type locality. “New York; Massachusetts; Maine.” Male, “New York”[M] [Very good condition, left hindwing margin partially damaged] Male, “New York”[M] [In fair condition, head missing] Female, “New York”[M] [In excellent condition, left antenna missing] Remarks: This material was found in the general collection and not labeled as a type. Poole (1989) states the type is in MSU. LaFontaine (1987) declared the type lost. The specimens before me agree with the original description and all were prob- ably in the original type series. I hereby designate the first male as the lectotype. A synonym of Euxoa declarata (Wlk.) (RWH 10755). Syneda deducta Morrison, 1874g[ 1875]: 220-221. Type locality. “Waco, Texas.” 2 females, “Texas” [?] [In very good condition] Remarks: Syntypes. These specimens were found in the general collection. There is also an unsexed specimen from “Tex” in the MCZ. Bulia (RWH 8614). Caradrina derosa Morrison, 1875b: 121. Type locality. “New Jersey. Received from Mr. W. V. Andrews.” Female, “N.J.”[T] “Type”[T], “Fr. Australia =/2fiocm capularis " \S] [The an- tennae, right legs and abdomen destroyed by dermestids] Remarks: The holotype; extralimital. A synonym of Rictonis capularis (Gn.) (Poole, 1989). 82 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Agrotis digna Morrison, 1875b:115. Type locality. “Texas.” Male, “21/9[?]”; “Texas” [T] [In very good condition] Female, 4/8.[?]”; “Texas” [T] [Right antenna missing, otherwise in good con- dition] Remarks: Status uncertain. This type was found unlabeled in the general collec- tion. Smith (1893) stated that the type, a single specimen, was in the Tepper collec- tion and another in the Peabody Academy of Sciences. Euagrotis (RWH 10908). Orthosia dijferta Morrison, 1875d:67. Type locality. “New York. Received from. . . Mr. Fred Tepper.” Male, “Type”[M]; "'dijferta U.S. Morr”[T] [A well marked specimen with all legs missing except the right metathoracic] Remarks: The holotype. Hardwick (1996) had no information regarding the lo- cation of this type. A synonym of Rhodoecia aurantiago (Gn.) (RWH 11065). Heliothis diminutivus Grote, 1873c: 148— 149. Type locality. “California (Mr. Hy. Edwards, No 204). Ten specimens examined.” Female, “California” [M]; “Type[C] [In good condition, right antenna missing] Female, “Cal.”[?] [In good condition, only right pro and meso legs present] Female, no labels. [Right pro and meso legs missing, in good condition] Remarks: Paralectotypes. Hardwick (1996) designated a specimen in the USNM as the Lectotype. Smith (1893: 291) stated that the “types are in the Tepper Collec- tion and in the British Museum.” Heliothodes diminutivus (RWH 11058). Lithophane disposita Morrison, 1874c: 109. Type locality. “Canada, Mass., New York. In April and May.” 3 males, “New York”[M] [All three in very good condition except one without an abdomen] Remarks: Syntypes. These specimens were found in the general collection and probably were part of the original type series of disposita (RWH 9892). Type(s) at MSU (Poole 1989). Demas diversicolor yiorrison, 1874b[1875]:132-133. Type locality. “New York. From my collection. Massachusetts, Sept. 16, 1874. (Mr. Roland Thaxter).” Male, “Tepper” [ty] ; “Massachusetts 232” [M] [Type in good condition with left forewing tom] Male, no data. [Left legs missing, well marked] Female, no data. [Legs and left antenna missing] Remarks: Only one type in the collection is labeled, and this is the MA specimen. The status of the two unlabeled specimens is not known. Smith (1893: 146) states that “the type is in the Tepper collection; another specimen is in the British Museum and is also marked type” while Poole (1989) states the type is at MSU. Lynnette Warshow (pers. comm. 1994) indicated 2 specimens from the 14th and 31st of Sep- tember from Newtonville MA with the label R[oland].T[haxter].C[ollection]. at the MCZ. It is not known whether types of this species are at the BMNH. The MCZ material is likely the better candidate for lectotype designation. Meropleon (RWH 9427). 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 83 Mamestra dodgei Morrison, 1875a:90-91. Type locality. “Nebraska (G.M. Dodge).” Male, “Nebr”[?]; ""Mamestra Dodgei, Type Mor”[M] [In very good condition, right antenna missing] Remarks: The holotype. A synonym of Lacinipolia lorea (Gn.) (10405). Heliaca dubitans Tepper, in Smith 1883:246.” Type locality. “Nevada.” Male, “Type”[C]; “Nevada” [T] [In excellent condition] Remarks: Holotype (Hardwick 1996). A synonym of Schinia perminuta (Hy. Edw.) (RWH 11091). Taeniocampa earina Morrison, 1874b[1875]: 158— 159. Type locality. “California.” Female, ""earina, Morr Type California 412”[M] [Some verdigris at the base of the pin, right antenna missing] Remarks: The holotype. The author of earina is not Harvey in (Hodges, 1983). A synonym of Lasionycta insolens (Grt.) (RWH 10366). Dicopis electilis Morrison, 1875b, 114-115. Type locality. “Easton, Penn. From Mr. W. H. Stultz.” Female, “Type”[C] [In poor condition with dermestid damage to abdomen and maculation somewhat rubbed] Remarks: The holotype (Poole, 1994). Psaphida (Poole, 1989, 1994) (RWH 10012). Agrotis exsertistigma Morrison, 1874b[1875]: 166. Type locality. “California.” Female, “So Cal”[T]; “Ty[pe]”[C] [Specimen in excellent condition] Remarks: This female was curated as a type of binominalis (see binominalis) but I have reason to believe that this is the Morrison type of exsertistigma, and curated under the Smith name. In Smith (1893) he states that “this is the exsertistigma of the Edwards collection, and like one specimen so labeled [exsertistigma] in Mr. Tepper’s collection.” Which may be interpreted as stating that the type of exsertis- tigma is to be found at MSU even though there was no type for that name curated in the collection. I would expect the type for exsertistigma to be at MSU since it is a Morrison name, and it is likely that the female above, curated with male types of binominalis, is in fact the type for exsertistigma; (RWH 11047); misspelled as ex- ertistigma in Hodges (1983). A synonym of Rhynchagrotus formalis (Crt.). Agrotis extranea Smith, 1888a:459. Type locality. “Montana. A single female specimen from Mr. Tepper.” Female, “Mt. Hood”[T]; “Type”[C]; ""Euxoa slide MSU No. 12.” [In very good condition, the prothoracic legs missing] Remarks: The holotype (Lafontaine, 1987). Euxoa (RWH 10708). Segetia fabrefacta Morrison, 1874b[1875]: 146-147. Type locality. “One specimen taken at Tuckemuck Island near Nantucket, and now in the collection of A. R. Grote. Another taken at Brooklyn, N.Y., in my possession.” 84 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Male, “Tepper”[T]; “New York 103“ [M]; “Type“[C] [The well marked spec- imen is in poor condition, the right wing has fallen off, the abdomen is lost to dermestid damage, and the legs and right antenna are missing.] Remarks: Syntype. The male above is likely the Morrison specimen mentioned in the original description. There is no specimen in the series labeled Nantucket or Tuckemuck Island, and since this was in Grote’s collection, it is presumed that this specimen is in the BMNH (Poole, 1989). A synonym of Platysenta sutor (Gn.) (RWH 9699). Lithophane fagina Morrison, 1874c: 1 15-1 16. Type locality. “Cambridge Mass., April 15, 1874. From my collection.” Male, “Type“[CJ; ""fagina Morr. Type. Camb. Ms 4,15 1874”[M] [The thorax severely damaged by verdigris and the right wings appear to have been glued to the thorax] Remarks: The holotype (RWH 9917). Agrotis fauna Morrison, 1876:237. Type locality. “Guadaloup island, lower Cal.” Male “Type”[C]; ""Euxoa slide MSU No. 15” [ty] [Type in poor condition, legs broken and damaged by dermestids, left antenna missing] Remarks: The holotype. Smith (1893: 98) states that the “type, a miserable spec- imen, is in the Tepper Collection.” Extralimital (Poole, 1989). Pachnobia ferruginoides Smith, 1890:56. Type locality. “Montana.” Male, “Montana” [T]; “Type”[C] [In good condition] Female, “Montana” [T]; “Type”[C] [Excellent condition] Remarks: Paralectotypes. Todd (1982) designated a USNM specimen as the lec- totype. A synonym of Paradiarsia littoralis (Pack.) (RWH 10992). Dryobota fibulata Morrison, 1874c: 112. Type locality. “Quebec. Can., in my collection from EX. Belanger.” Female, ""Dryobota fibulata Morr Can Type”[M,T] [Abdomen and antennae absent, markings prominent] Remarks: The holotype. The specimen is unusual in that Morrison departs from his trademark triangular label and uses a square label. The words ""Dryobota fibulata Morr Can” appear to be in Morrison’s hand, and a portion of the “Q” in Quebec appears to have been cut off the label. “Type,” on the other hand, is written in red ink in Tepper’s hand. A synonym of Eremobina claudens (Wlk.) (RWH 9396). Segetia fidicularia Morrison, 1874b[1875]:145. Type locality. “Adirondack Region, N. Y. A single specimen in the collection of my friend Mr. F. C. Bowditch, of Harvard Law School.” Female, “Milford N. H. 94” [M]; ""Segetia fidicularia Morr. 94” [M] [Right antenna, left pro and meso and right meso and meta legs missing, mold damage] Remarks: This specimen was found, unlabeled in the general collection. The lo- cality of this specimen does not match the original description; it is not known if this is the type. Neither Smith (1893) or Poole (1989) indicated the location of the type. It is, however, possible that the locality discrepancy is an error in the original 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 85 description. In the absence of other convincing type material, this specimen with a determination by Morrison, may be a good candidate as a type. A synonym of Caradrina multifera (Wlk.) (Poole, 1989) (RWH 9657). Heliophila flabilis Grote, 1881a: 15. Type locality. “Long Island, near the sea shore, in May; Mr. Tepper.” Male, “L. I. May 20 [18]77,’’[T]; “Type“[C] [Type in fair condition; both meso legs and right antenna missing] Remarks: Syntype. Smith (1893 p. 189) states the “types are in the Tepper collection and in the British Museum. The species was described from material taken by Mr. Tepper.” Poole (1989) states that the type is in the BMNH. A synonym of Leucania extincta (Gn.) (Poole, 1989) or subspecies of L. extincta flabilis (RWH 10439a). Agrotis flavicollis Smith, 1888a:456. Type locality. “Montana. One female specimen from Mr. Tepper’s collection.” Female, “Montana” [T]; “Tepper” [ty]; “Type”[C]; “Euxoa slide MSU No. 13”[ty] [In good condition, but only the right meso and left meta leg present] Remarks: The holotype (Todd, 1982). Euxoa (RWH 10864). Homohadena fortis Grote, 1880a:257. Type locality. “Nevada.” Female, “Nevada” [T]; “Type”[C] [In very good condition, the left pro and right pro and meso legs and right antenna missing] Remarks: Status uncertain. Poole (1989) states that the types are in the BMNH. Smith (1893) stated the type was in the Tepper collection. In the absence of other material, this is probably the holotype. Homoncocnemis (RWH 10068). Ablepharon fumosum Morrison, 1874a:275. Type locality. “Massachusetts. May 24th and 26th. Coll. H. K. Morrison.” Male, “Cambridge, Ms 5/21 288” [M]; “Type”[C] [In excellent condition] Male, “Cambridge, Ms 5/30 288” [M]; “Type”[C] [In excellent condition] Remarks: Status uncertain. Although the precise day of May is not in agreement with the original description, it is difficult to conclude that these are not syntypes. A synonym of Simyra henrici (Grt.) (RWH 9280). Hadena fuscimacula Grote, 188 lb: 262. Type locality. “No locality given.” Male, “Florida” [T]; “Type”[C] [In very good condition] Female, “Florida” [T]; “Type”[C] [In good condition, left antenna missing] Remarks: Poole states that the types are in the BMNH. It is not known if this specimen is also a part of the original type series. Elaphria fuscimacula (Grote) (RWH 9675). Homoptera galbanata Morrison, 1875c[1876]:435-436. Type locality. “Glencoe, Nebraska. From Mr. G. M. Dodge (No. 48).” Male, “48”; “Type”[C] [In excellent condition] Remarks: Probably the holotype. Zale (RWH 8692). Calocampa germana Morrison, 1874e:192. Type locality. “Adirondack Mountains. (Mr. Bowditch).” 86 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) 2 males, 1 female, “New York”[?] [In very good condition] Remarks: Syntypes. These specimens were found in the general collection without type labels. Poole (1989) states that the type is at MSU. Smith (1893) did not indicate the location of the type. The handwriting of the three specimens is not known, but may be that of Bowditch. A male from the above series has been selected and labeled the lectotype. A subspecies of Lithomoia solidaginis (RWH 9878a). Agrotis gladiaria Morrison, 1874b[ 1875]: 162-163. Type locality. “Massachusetts.” Male, “Cambridge Mass. 177” [M]; “Type”[C] [Right antenna missing, oth- erwise in very good condition] Remarks: The original number of types at the time of description is not known. If no other type material of this species is found, this is probably the holotype. There may be additional material for this species at other collections (RWH 10648). Melicleptria graefiana Tepper, 1883:245. Type locality. “Southern California.” Male, “So Cal”[T] [In fair condition, only left pro leg present, left antenna broken] Male, “So Cal”[?] [In good condition, the right antenna missing] Female, “So Cal”[T] [Only left meso leg is present] Remarks: The specimens above were curated as types for this species but are not, the holotype is at the USNM (Hardwick, 1996). Hardwick (1958) acknowledged the above specimens as Schinia pulchripennis (Grt.). Since California is the type locality for Schinia pulchripennis, there is a strong chance that the type(s) above represent the type series for pulchripennis, types which Hardwick (1996) presumed lost. Valeria grotei Morrison, 1874a: 274. Type locality. “Massachusetts. Specimens taken in Cambridge from April 10th to 26th, coll. H. K. Morrison.” Male, ""grotei, Morr. Type, Cambridge M[a]”, April 14, 331”[M]; “Type”[C] [A well marked specimen in good condition; left pro and meso legs and antenna miss- ing] Remarks: The holotype (Poole, 1994). The location of the other specimens indi- cated in the description is not known, nor does Poole (1994) mention these other specimens. Copivaleria (RWH 10021). Schinia hulstia Tepper, in Smith 1883:228. Type locality. “1 female Texas? Type, Coll. Tepper.” Female, “Texas” [?]; “Type”[C] [In fair condition] Remarks: The holotype (Hardwick, 1996) (RWH 11193). Mamestra illabefacta Morrison, 1874c: 141-142. Type locality. “Beverly, Mass. June 26, 1869, Edward Burgess.” Male, ""illabefacta Morr. Type Beverly, Mass 18”[M] [In very good condition, the left pro and meta leg and left antenna missing] Remarks: Smith (1893: 123) stated that there is a type “in the Tepper collection, and another ... in the British Museum.” McCabe (1980) did not indicate the location of the type. Given all the trademarks of Morrison, and despite the discrepancy in the date of capture, this is likely the holotype. A synonym of Lacanobia lilacina (Harvey) (RWH 10307); Trichordestra (Poole, 1989). Misspelled as illebefacta in Hodges (1983). 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 87 Acronycta increta Morrison, 1874c; 131. Type locality. “New York. Several specimens received from Messrs. Fred Tepper and E. L. Graef. Types in the collections of the Buffalo Society of Natural Sciences, and H. K. Morrison.” Female, “Brooklyn N. Y.”[M]; “Type”[T] [The well marked specimen has the right wing missing] Remarks: Syntype. The location of the other specimens is not known. Acronicta (RWH 9249). Laphygma inflexa Morrison, 1875d:65-66. Type locality. “Jacksonville, Fla.” Female, “Florida” [T]; “[illegible]”; “Type”[C]; Lyssa (Laphygma) filifera Wlk.”[Sm] [In good condition] Remarks: The holotype. A synonym of Collomena filifera (Wlk.) (RWH 8982). Agrotis infracta Morrison, 1875b: 115. Type locality. “Colorado (T. L. Mead); Texas (Belfrage).” Female, “Col”[T]; “Type”[C]; ''Euxoa slide MSU No. 3”[ty] [In fair condi- tion] Male, “Col”[ty]; “male. ’’[symbol] [In poor condition, the antennae are missing and the right valve missing] Remarks: The male was found in the general collection. Lafontaine (1987) stated that he had designated the female as the lectotype, but no such label was found on the specimen. I have since added a lectotype label. Euxoa (RWH 10850). Hadena inordinata Morrison, 1875d:63. Type locality. “Massachusetts, in June from our collections.” Female, “Mass”[M]; “Newtonville June 16, 1874”[M]; “Type”[C] [In very good condition] Remarks: The original “from our collections” hints that there were a number of specimens. The absence of an explicit indication of the name inordinata is not typical of Morrison. In the absence of more convincing material, this may be a good can- didate for lectotype designation. This is certainly a type if no other types exist. Apamea (RWH 9353). Taeniocampa intractata Morrison, 1874b[l 875]: 160-161. Type locality. “Habitat, St. Louis, Mo., Prof. C. V. Riley.” Male, "'intractata, Morr Type, Cen. Missouri 414”[M]; “Type”[C] [In good condition, antennae missing] Female, no data. [Right hindwing and antennae missing] Male, no data. [Extensive dermestid damage to abdomen and thorax] Remarks: Syntypes. The status of the male and female without data is not known. The labeled male is undoubtedly a valid type, however, Poole (1989) also indicates that a type is at the USNM. Himella fidelis (RWH 10502). Oncocnemis tricolor Smith, 1888b: 19. Type locality. “Colorado. Three specimens, all females, 1 Coll. U.S. National Museum (Smith Coll.) the others Coll. F. Tepper.” Female, “Col”[T]; “Type”[C] [In fair condition, some dermestid damage to the abdomen, left pro leg missing] 88 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Female, “Col”[T]; “Type”[C] [Antennae missing, well marked] Remarks: The females above are paralectotypes. Todd (1982) designated a spec- imen in the USNM as the lectotype (RWH 10092). Plusia laticlavia Morrison, 1875f:98. Type locality. “New York. July 10, 1872. Described from a single specimen in good condition received from Mr. Fred Tepper. ” Male, “L. I. July 10, 1872’’[T]; “Type Morr”[M]; “laticlavia” [?] [In very good condition, right meta leg missing] Remarks: The holotype. There were fully three different plusiine species in this “type” series. From the original description, however, it is clear that Morrison had but a single specimen at the time of description. A synonym of Ar gyro gramma basigera (Wlk.) (RWH 8886). Panthea leucomelana Morrison, 1875c[1876]:428. Type locality. “Maine.” Male, “Maine” [T] [Both metathoracic legs missing] Remarks: This specimen was found in the general collection without a type label. This is very likely the holotype. Poole (1989) states the type is at MSU. A synonym of Panthea acroncytoides (Wlk.) (RWH 9177). Luceria loculata Morrison, 1874c: 110. Type locality. “Mass., New York. Four specimens examined and one of var. con- spicua^ Male, 'doculata, Morr Type Boston, MS, 44” [M] [A well marked specimen with hollow abdomen, genitalia appear intact] 2 males, no data. [Well marked but with some verdigris on thorax] Remarks: Syntypes. The location of the additional type material, or the status of the two unlabeled males is not known. A synonym of Luperina passer (Gn.) (RWH 9391). Luceria loculata var. conspicua Morrison, 1874c: 111. Type locality. “Mass., New York. Four specimens examined and one of the var. conspicua y Male, “Boston Ms 59”; “Type”[M] [In good condition, extensive damage to abdomen and genitalia] Remarks: One specimen is indicated and presumed to be the holotype. The type locality of conspicua is not clearly indicated in the original description. A synonym of Luperina passer (Gn.) (RWH 9391). Heliothis lucens Morrison, 1875d:69. Type locality. “Mass., Nebraska.” 2 males, 3 females, “Nebr.”[M] [In very good condition] Remarks: Paralectotypes (Hardwick 1996). These five specimens were in the gen- eral collection and not labeled as types. Only one label appears to be in Tepper’s hand, the other in Morrison’s. Schinia (RWH 11174). Penthetria majuscula Henry Edwards, 1881a:80. Type locality. “Georgia. H. K. Morrison. Type. Coll Hy. Edwards. Female. Type. F. Tepper.” 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 89 Female, “Ga”[T]; “female” [?]; “Type”[C] [In fair condition, the antennae missing] Remarks: Syntype. The male syntype is at the AMNH (Poole, 1989). A synonym of Cydosia aurivitta (RWH 8999). Agrotis manifesta Morrison, 1875d:116. Type locality. “Described from specimens in the collection of Mr. Fred Tepper. Hab. New York. In May.” Female, “Flatb[ush] May 25/72” [T]; “Type”[T] [In very good condition] Male, “Flatb[ush] May 25/72” [T]; “Type”[C] [In poor condition, dermestid damage to head, thorax, and abdomen] Remarks: Syntypes. Lectotypes have not been designated due to the poor quality of this material. Hodges (RWH 10666). Agrotis manifestolabes Morrison, 1874b[ 1875]: 166. Type locality. “Massachusettes, in the early spring.” Male, '"manifestolabes Morr. Type. Cambridge, April 29, 187[2] 183” [M] [In fair condition, the right legs and left antenna missing] Remarks: The holotype. A synonym of Cerastis tenebrifera (Wlk.) (RWH 10994). Oncocnemis meadiana Morrison, 1875d:60— 61. Type locality. “Colorado, Aug. 18 (No. 47, Mr. T. L. Mead).” Female, “Colo Aug, 18, 1872 216”[M]; “Type”[C] [Specimen in good condition] Remarks: The holotype (RWH 10098). Caradrina meralis Morrison, 1875g:215. Type locality. “Maine. Collection H. K. Morrison.” Male, “Maine” [M]; “Type”[C] [In excellent condition]. Remarks: The original extent of the type series is not known. This is probably the holotype. Caradrina Poole (1989)(RWH 9654). Carneades messoria var. confracta Smith, 1890:170. Type locality. “One form. . . has been named var. confracta. by Mr. Morrison in Mr. Tepper’s collection (Smith 1890).” Extracted from Todd (1982). Female, “Agrotis Messoria var. Confracta Morr. U.S.”[T]; "Euxoa slide MSU No. 8.”[ty] [Type in fair condition, right antenna missing] Remarks: The holotype (Todd, 1982). The name confracta, discussed by Smith (1890) was a manuscript name. The mention of a MSS name is attributed to Smith who formally published a discussion of a name which constitutes a valid description. A synonym of Euxoa pleuritica (RWH 10727). Zanclognatha minoralis Smith, 1895:34, Plate 2, fig. 6, plate 10 figs. 28, 29. Type locality. “Long Island, New York, probably.” Male, " Me gachyta minoralis Smith, Type”; “6” "CQCioiypQ, Zanclognatha mi- noralis by Todd” [Right palpus missing, otherwise in good condition] Remarks: This specimen was found by J. E. Zablotny in a slide box in a drawer of unsorted bees in the general collection. The lectotype of minoralis (RWH 8343) was designated by Todd (1982). Orthosia minuscula Morrison, 1874b[ 1875]: 147. Type locality. “Tuckernuck Island, near Nantucket. Mr. Edward Burgess.” 90 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Female, “Tepper”[ty]; “Nantucket 46“ [M]; “Type“[C] [In fair condition, an- tennae missing and markings somewhat rubbed] Remarks: It appears he had but a single specimen when he described it since he indicated displeasure in the “badly rubbed” specimen and hoped that “perhaps the discovery of a better specimen will show that this species should be referred to a different genus.” It is not known which, if either, is the holotype. Oligia (RWH 9416). Carneades misturata Smith, 1890:156. Type locality. “Colorado.” Male, “Col”[T]; “Type”[C]; '^Euxoa Slide MSU No. 7”[ty] [Left meso leg missing, antennae missing] Remarks: Todd (1982) stated that the type “in the Michigan State University collection has been selected, labeled, and is now designated the lectotype.” There is no lectotype label affixed to this specimen. I have since added a label. Euxoa (RWH 10766) of Morrison manuscript name. Taeniocampa modifica Morrison, 1874b[1875]:150-151. Type locality. “Massachusetts. A male from my collection captured July 16, 1874. A female from the collection of Mr. F. C. Bowditch, kindly lent me for deter- mination.” Male, “July 14, 1874 Cambridge, Ms 414” [M]; “Type”[C] [The left antenna missing] Male, [no labels, in very good condition] Remarks: The labeled male is probably the holotype. The male type was obviously collected by Morrison and is likely the specimen he mentions despite the discrepancy in the date on the specimen and that of the original description. The location of the Bowditch female is not known. The status of the unlabeled male is unknown. A synonym of Ulolonche culea (Gn.) (RWH 10567). Agrotis montana Morrison, 1875T94-95. Type locality. “Mountains of Colorado, above 12,000 feet, July 22 to Aug. 12. One specimen in the possession of Dr. A. S. Packard, Jr., and another in our own collection.” Female, “Col”[T]; ''montana Morr. Type Colorado above 12,000 ft 172”[M] [Dorsally flattened, abdomen eaten by dermestids, in extremely poor condition] Female, “Col”[T] [In fair condition] Remarks: Syntype. Morrison does not list the sex of the two specimens he had before him at the time of the description; the first female is probably a type. Smith (1893: 141) states that “Mr. Morrison’s type is in the Tepper collection, and is from the east.” Poole (1989) and Lafontaine (1987) state the type is at the MCZ. Lynette Warshow (pers. comm. 1995) stated the type(s) are at MCZ, however, there is no information associated with this entry on the Internet at this time. The presence of the trademark Morrison label suggests that the female, labeled montana by Morrison is also a (syn)type and probably refers to the “in our own collection” specimen. Euxoa (RWH 10856). Eutricopis nexilis Morrison, 1875f:102-103. Type locality. “Colorado, June 18.” 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 91 Male, “Colorado” [M]; “Type”[C] [In good condition, antennae missing] Remarks: Paralectotype. Hardwick (1996) designated a lectotype, now in the USNM. The locality label is one of Morrison’s and is in his handwriting, but he does not indicate the specific name on the label, which is unusual for a Morrison type (RWH 11062). Tarache obatra Morrison, 1875b: 124 Type locality. “Louisiana.” Female, “Louisian[a]”[T]; “Type”[C] [In very good condition, antennae miss- ing] Remarks: The holotype. Spragueia (RWH 9130). Eustrotia obaurata Morrison, 1874b[ 1875]: 154. Type locality. “Massachusetts.” Female, ^'obaurata, Morr Type. Mass 250” [M] [In fair condition, only the left meta leg complete] Remarks: The original number of specimens in the type series is not known; probably the holotype. A synonym of Nola pustulata (Wlk.) (RWH 8989). Mamestra olivacea Morrison, 1874b[1875]:143. Type locality. “New York; New Hampshire.” Female, “N. Y. State” [M] [In good condition, the abdomen is broken off, and is associated with the specimen in a genitalia vial] Remarks: This specimen was found in the general collection and not labeled as a type. Six other specimens were associated with it, all in Tepper’s hand, and from western North America. The location of the material from NH is not known. If the NH material cannot be found, this is probably the holotype. Poole (1989) states type is at MSU. Lacinipolia (RWH 10406). Agrotis Olivia Morrison, 1876:238. Type locality. “Utah (T.L. Mead).” Male, “Utah 186”[M]; “Type”[C] [In good condition, only the right pro leg present] Remarks: The holotype (Lafontaine, 1987). Euxoa (RWH 10741). Agrotis opipara Morrison, 1874b[ 1875]: 165-166. Type locality. “The alpine region of Mt. Washington.” Female, “Mt. Washington, July 11, 1874.”[ty] [In good condition] Remarks: This specimen was found in the general collection, is unlabeled and is likely a type. Lafontaine (1987) stated that the type is in the BMNH. The MSU specimen also appears to be a syntype. A synonym of Euxoa dissona (Mosch.) (RWH 10706). Segetia orbica Morrison, 1874g[1875]:216-217. Type locality. “Waco, Texas.” Female, "'orbica Morr. Type. Waco. Tex. 7 28 125” [M]; “Tepper”[ty]; “Type”[C] [In good condition, the right pro and both meso legs and the left meso leg and left antenna are missing] Male, “19”[?] [In good condition, the antennae are missing] Remarks: Another putative type of orbica is at the MCZ (L. Warshow, pers. 92 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) comm. 1995). The unsexed specimen is at the MCZ, without specific label data and is not likely a type. The male is of uncertain validity, so the female above is selected and designated as the lectotype. A synonym of Amyna octo (Gn.) (RWH 9070). Agrotis orbicularis Smith, 1888a:460. Type locality. “Colorado (Bruce). A single female is in Mr. Tepper’s collection.” Female, “Nevada”[T]; “Type”[C]; “Euxoa slide MSU No. 14”[ty] [In very good condition] Remarks: Despite the discrepancy in label data, Lafontaine (1987) recognized this specimen as the holotype. A synonym of Euxoa cooki McD. (RWH 10858). Heliophila oxygala Grote, 1881a: 14. Type locality. “One specimen in Mr. Tepper’s collection, one male in my own.” Female, “Col.”[T]; “Type”[T] [The well marked specimen is in poor condition, the left pro leg the right antenna is missing, and glue appears to have been applied to the left wings and the right hindwing. Thorax damaged by dermestids] Remarks: Syntype. Poole recognized types in both BMNH and MSU. Smith (1893 p. 185) states that the “type is in the British Museum. I have a specimen carefully compared with it. Another specimen, also marked type, is in the Tepper collection.” The red inked labels “Col.” and “Type” are in Tepper’s hand. Aletia (RWH 10436). Hadena paginata Morrison, 1875d:64-65. Type locality. “Florida (Mr. C. J. Maynard).” Male, “Florida”[T]; “Tepper”[tyJ; “Type”[C] [In poor condition; the abdomen destroyed by dermestids] Remarks: Probably the holotype. A synonym of Elaphria nucicolora (Gn.) (RWH 9676). Aedophron pallens Tepper, In Smith 1883:215. Type locality. “Southern California.” Remarks: Type presumed lost. This species was described from the Tepper col- lection in a paper by Smith. I have been unable to find a specimen of this species in the collection. Nocloa (RWH 9795). Agrotis pallipennis Smith, 1888a:46 1-462. Type locality. “Colorado. Specimens are with Messrs. Hulst, Graef, and Tepper.” Female, “Col.”[T]; “Tepper” [ty]; “Type”[C] [In good condition] Male, “Col”[T] [In fair condition] Remarks: Paralectotypes. Todd (1982) mentions only one specimen at MSU and makes no mention of the male above. In addition, he stated that the female MSU specimen was in poor condition and that the abdomen appeared glued to the thorax. I have been unable to find any indication that the abdomen was glued. A lectotype of this species was designated from the U.S.N.M. (Todd 1982). Euxoa (RWH 10765). Heliophila patricia Grote, 1880c:46. Type locality. “Colorado. Coll, of Mr. Tepper.” Female, “Cal”[T]; ""Heliophila patricia Grote Type”[G] [In excellent condition] Remarks: The holotype. Neleucania (RWH 10610). 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 93 Tarache patula Morrison, 1875d;69— 70. Type locality. Texas. September 11. Expanse 15mm.” Male, “Type”[M]; Texas, Sept. 11. 263” [M]; “Type”[C] [In good condition] Remarks: Two female specimens from Texas were also in the type series. Both specimens have handwritten labels, apparently written by two different individuals — neither appears to be in Morrison’s hand; they are not likely types but have not been removed from the type series. The male, is selected and designated as the lectotype. A synonym of Eublemma recta (Gn.) (Poole, 1989); Eumestleta recta (Gn.) (RWH 9078). Mamestra passa Morrison, 1874b[ 1875]: 139— 140. Type locality. “California. From my collection.” Female, “Cal”[M]; “Type”[C] [Right meso, left meso and meta legs and an- tennae missing] Remarks: Graphania (Poole 1989), extralimital. Lower California (Baja). Syneda pavitensis Morrison, 1874f[ 1875]: 221. Type locality. “Waco, Texas.” Female ''pavitensis, Morr. Type. Sept. 7 Waco. Tex. 368” [M] [In good to fair condition, the right meso leg missing, and verdigris building around the base of the pin] Remarks: Syntype. Smith (1893: 325) states that the “type[s] are in the Museum of Comparative Zoology at Cambridge and have been correctly referred as sexes of the same species.” A duplicate type of pavitensis is in the Tepper collection.” No indication as to the number of specimens in the original type series. A synonym of Bulia deducta (Morrison) (RWH 8614). Heliothis pauxillus Grote, 1873b:118, pi. 3 fig. 6. Type locality. “Colorado Territory, (Coll. Theo. L. Mead No. 9).” Male, "pauxillus Gr Type Colo July 7 300” [M] [Well marked specimen in fair condition, antennae missing, right meso and meta and left meso legs missing] Remarks: The male is unusual is that it is a Grote name in Morrison’s handwriting. The specimen is the holotype (Hardwick, 1996). A synonym of Schinia villosa (Grt.) (RWH 11083). The year of publication (1875) is in error for pauxillus in Hodges (1983). Lithacodia penita Morrison, 1875d:71. Type locality. “New York. One female specimen received from Mr. Fred Tepper.” Female, “Type”[C]; [Specimen in poor condition, verdigris expanding in tho- rax, left antenna missing] Remarks: The holotype. This is the presumed type of penita, however, labels that would associate the specimen with Tepper are not present. Poole (1989) indicates the type is at MSU. A synonym of Spartiniphaga includens (Wlk.) (RWH 9434). Morrisonia peracuta Morrison, 1874c: 114. Type locality. “Doubtful, probably Texas, perhaps California. In the collection(s) of H. K. Morrison.” Male, “Cal”[M]; “Type”[C] [The left antenna missing, otherwise the type in very good condition] 94 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Remarks: The holotype. The locality information on this specimen is incorrect; it is an Australian species, a synonym of Persectania ewingii Westwood (Poole, 1989). Bryophila percara Morrison, 1874g[1875]:213-214. Type locality. “Waco Texas.” Female, “Texas” [T]; “Type”[C] [In very good condition] 3 Males, “Tex”[ty] [In good condition] Remarks: Syntypes. It is not clear how many specimens were in the type series; there can be little doubt that the female is from the original type series, the males, however are more problematic. Each specimen is labeled with a typewritten “Tex.” on white paper compared to the female which is in Tepper’s hand and written in red ink. I retain the three specimens, but I suspect they were added at a later date. There are also syntypes of this species at MCZ (L. Warshow, pers. comm. 1995). Emar- ginea (RWH 9718). Scotogramma perplexa Smith, 1888a:469. Type locality. “Colorado. The type is a unique female in the collection of Mr. Tepper.” Female, “Col”[T]; “Type”[C] [In fair condition] Remarks: The holotype (Todd, 1982). Lasionycta (RWH 10352). Agrotis perpolita Morrison, 1876:237. Type locality. “Orono Maine.” Male, “Maine” [M]; “Type”[C]; ''Euxoa Slide MSU No. 4”[ty] [In good con- dition] Remarks: The holotype (Lafontaine, 1987). Euxoa (RWH 10865). Orthosia perpura Morrison, 1875d:66. Type locality. “New York” Female, “perpura, Morr Type Colo. July 20 495” [M] [Left meso and meta and right pro and meso legs and right antenna missing] Remarks: The holotype. A synonym of Lasiestra impingens (RWH 10339). Heliothis persimilis Grote, 1873b: 1 17-1 18, pi. 3, fig. 11. Type locality. “Colorado Territory (coll. Theo. L. Mead, No. 6).” Female, “Colo”[M]; “Type”[C] [Type in fair condition, the right meso and meta legs, and left meso leg missing] Remarks: The holotype (Hardwick 1996). Schinia (RWH 11103). Agrotis personata Morrison, 1876:238-239. Type locality, ''personata comes only from Central Illinois.” Female, “Ohio”[M], A. personata, Morr. Type Ohio”[M]; “Euxoa Slide MSU No. 2”[ty] [Left hindwing missing, right forewing chipped at the apex] Remarks: Despite the disagreement in the type locality, this is the holotype (La- fontaine 1987). A synonym of Euxoa detersa (Wlk.) (RWH 10838). Hodges (1983) is in error for personata as a Grote name. Agrotis plagigera Morrison, 1874b[ 1875]: 163. Type locality. “Colorado. Expanse 33 mm.” Female, “Tepper” [ty] [In good condition, right antenna missing] 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 95 Remarks: This specimen was found unlabeled in the general collection. In the absence of any probable type specimens at the time of his study, Lafontaine (1987) declared the type lost. This is a likely candidate for type, despite the fact that no locality information is associated with the specimen. The original accession notes of the Tepper collection states that the type of this species was in this collection; evi- dently Cook did not label it as a type. Euxoa (RWH 10804). Agrotis proclivis Smith, 1888a:45 3-454. Type locality. “Arizona. One male (Tepper) and One female (Neum.).’’ Male, “Arizona” [T]; “Type”[C] [In poor condition, the head, thorax and ab- domen is severely damaged by dermestids and the right valve of the genitalia broken] Remarks: Todd (1982) designated the female in the MSU as the lectotype. No such label is found on the specimen, it has since been added. Loxagrotis (RWH 10871). Mamestra promulsa Morrison, 1875L97. Type locality. “Colorado (Mr. T. L. Mead), July 20.” Female, “Col. July 219” [M]; “Type”[C] [Left pro meso, and right pro, meta legs missing, somewhat rubbed and dorsoventrally depressed] Female, “Col”[T]; “Type”[C] [Legs absent, right antenna missing] Remarks: Syntypes. Lasionycta (McCabe, 1997, pers. comm.) (RWH 10344). Acronycta pudorata Morrison, 1875L93. Type locality. “New York; Canada. Specimens received from Messrs. Fred. Tepper and FX. Belanger.” Male, “[Qu]ebec Can”[M]; ""pudorata, Morr Type”[M] [In fair condition, the right legs missing and the right forewing tom off at the apex] Remarks: Syntype. The male, is clearly a Morrison type, and is unusual in having two of the trademark Morrison labels. No Cook (rhomboidal) type label is associated with these specimens. Smith (1893: 31) gives Tepper collection as location of the (holo)type. Poole (1989) gives the BMNH as the location of the type. A synonym of Acronicta grisea (Wlk.) (RWH 9212). Mamestra quadrannulata Morrison, 1875c[1876]:430-431. Type locality. “Glencoe, Dodge Co., Nebraska. From Mr. G. M. Dodge (No. 38).” Male, “38”[?]; “Tepper” [ty]; “Type”[C]; [A well marked specimen, left valves and tegumen nearly completely destroyed] Remarks: This specimen lacks the trademark Morrison label or any handwriting resembling Morrison’s, but matches the original description in having the number 38. This is likely a type. Faronta (RWH 10430). Hadena rasilis Morrison, 1874b[ 1875]: 158. Type locality. “St. Louis. From my collection; received through the kindness of Mr. C. V. Riley, State Entomologist.” Female, ""rasilis, Morr Type. St Louis, Mo 34” [M] [In good condition, the abdomen missing] Remarks: The holotype of rasilis. A synonym of Elaphria grata Hub. (RWH 9684). 96 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Agrotis redimicula Morrison, I874b[1875]: 165. Type locality. “Colorado (T. L. Mead); Albany N. Y. (J. A. Lintner):Massachu- setts.” Female, “Cambridge Mass’’[?]; “Type”[CJ; ""Euxoa Slide MSU 6.”[ty] [A very well marked specimen; both antennae broken near head] Remarks: The label “Cambridge Mass,” is in an unfamiliar hand. Lafontaine (1974) designated the MSU specimen as the lectotype, but no such label is associated with this specimen; I have since added a lectotype label. Euxoa redimicula (RWH 10851). Homohadena retroversa Morrison, 1874b[1875]: 157-158. Type locality. “Central Missouri. Prof. C. V. Riley.” Male, “Mo”[MJ; “June 22, 1872” [M]; “Type”[C] [In good condition, anten- nae missing] Remarks: Smith (1893) states that type is in the Tepper collection and a duplicate in the USNM. Poole (1989) states that the type is at MSU. A synonym of H. infixa (Wlk.) (RWH 10065). Agrotis rileyana Morrison, 1874b[ 1875]: 166. Type locality. “St. Louis, Mo.” Male, “Mo”[M]; “Missouri 188”[M]; “Type”[C] [In good condition] Female, “Mo”[M] [In very good condition] Remarks: Possibly the holotype; the original number of specimens is not known. Onychagrotis (RWH 10669). Oncocnemis riparia Morrison, 1875e:213. Type locality. “Mr. Fred Tepper has just sent me a pair of insects taken on the shore of Long Island.” Male, “L. I. Sound, July 7/75” [T] [In very good condition] Female, “Tepper” [ty] [Meso and meta legs and antennae missing] Remarks: Syntype(s). The types of this species were found in the general collec- tion and probably are types. Smith (1893) stated the type was in the BM and the Tepper collection. Poole (1989) gives MSU as the type location. The lectotype should not be designated until the BMNH can be checked for types of this name (RWH 10135). Eutolype rolandi Grote, 1874c: 198-199. Type locality. “Massachusetts, April 8th to 15th (Mr. Roland Thaxter, No. 1019); Missouri, April (Prof. C. V. Riley).” Male, “April 15th 1874 Nville Mass R. Thaxter S[?]p. No. 1019” [In very good condition] Remarks: Paralectotype. This specimen was found in the general collection and was not labeled as a type. It is clearly part of the original series, with a label that is presumably in Thaxter’s hand. Poole (1994) designated a specimen in the BMNH with Grote’s type label as the lectotype. This is also a paralectotype. Psaphida (RWH 10014). Mamestra rufula Morrison, 1875d:62. Type locality. “Massachusetts, New York, Illinois, Missouri, Indiana.” 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 97 Male, “G. L, July 4/74” [T] [In poor condition, antennae and abdomen missing] Remarks: The holotype. This type was found in the general collection. McCabe (1980) was unable to find the type in other collections. Poole (1989) indicated MSU as the location of the type. A synonym of Spiramater lutra (Gn.) (Poole 1989) RWH 10301. Mamestra rugosa Morrison, 1875b: 119. Type locality. “Maine. From Prof. C. H. Fernald, of Orono.” Male “Maine” [M] [Abdomen partly destroyed by dermestids, the valves partly broken] Remarks: This specimen was found in the general collection and not labeled as the lectotype. In his revision of the Folia complex, McCabe (1980), designated this specimen as the lectotype. However, no such label could be found on this specimen; a label has since been added. Lacanobia [Trichordestra] (RWH 10302). Agrotis scropulana Morrison, 1874b[1875]:165. Type locality. “The alpine region of Mount Washington.” Male, “Mt. Washington July 18, 1874”[ty] [In excellent condition] Remarks: The holotype. Xestia (Poole, 1989) type at MSU. Probably the holotype. Pachnobia (RWH 10938). Hydroecia semiaperta Morrison, 1874d:105. Type locality. “Mass., New York. Not uncommon. Coll, of H. K. Morrison.” Female, ''semiaperta, Morr. Type. Beverly, Ms 7.8.72 107” [M] [In excellent condition] Remarks: Syntype. The location of the other specimen(s) is not known. A syn- onym of Tricholita signata (Wlk.) (RWH 10627). Lithophane semiusta Grote, 1873a:34. Type locality. “Male and female, (Mr. J. Pettit).” Male, “Type”[C]; "semiusta Grote Type Eastern State 77” [M] [In good con- dition, the right antenna broken near base] Female, “N. Y. State” [M] [In poor condition, abdomen missing] Male, “N. Y. State” [M] [In excellent condition] Remarks: The status of the male marked type by Morrison is not known. The specimens marked “N Y State” are probably not types, but could possibly be types since “Eastern State” is ambiguous. Poole (1989) states the type is in the BMNH while Smith (1893 p. 228) stated that the “type is in the BMNH and another in the Tepper collection.” (RWH 9885). Glaea sericea Morrison, 1874b[ 1875]: 151. Type locality. “Boston Mass.” Female, “Boston Ms 114”[M]; “Type”[C] [In excellent condition] Remarks: It is not known whether types are in other institutions. Poole (1989) states that the type is in MSU. Probably the holotype. Chaetaglaea (RWH 9950). Agrotis serricornis Smith, 1888a:458. Type locality. “Southern California.” Male, “So Cal”[T]; “Tepper”[ty]; “Type”[C]; “Euxoa Slide MSU 10”[ty] [In 98 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) poor condition, the right hindwing partly destroyed, the right antenna missing and the valves partly destroyed] Remarks: Todd (1982) concluded that this was the holotype by the absence of any other specimens labeled “Type.” Euxoa (RWH 10744). Agrotis Simplicius Morrison, 1874b[ 1875]: 164. Type locality. “Texas.” Female, “Texas.” [M] [The specimen is in very good condition] Remarks: Syntype. This specimen was found in the general collection and is prob- ably a type. Lynette Warshow (pers. comm. 1995) indicates their unsexed specimen as the holotype. It is likely that both are syntypes, and the MCZ specimen is not the holotype. Poole (1989) stated that the type is at MSU. Euagrotis (RWH 10907). Agrotis stigmosa Morrison, 1874b[1875]:163. Type locality. “Mass. N. Y.” [Morrison] Male, ''stigmosa Morr Type, Camb. MS. June, 166” [M] [Abdomen missing, otherwise in good condition] Male, “Mass.”[M] [In fair condition, the abdomen is missing] Remarks: The second male specimen was found in the general collection, and is also likely a type. Poole (1989) stated that the type is at MSU. There are no speci- mens of this name at MCZ. I hereby designate the male, marked and labeled “stig- mosa,” as the lectotype. RWH (10658). Hadena stipata Morrison, 1875d:64. Type locality. “Illinois. One specimen received from Thos. E. Bean.” Female, “Illinois” [?]; “Type”[C]; Type in very poor condition, abdomen miss- ing, thorax destroyed by dermestids, wings tattered and worn, head missing] Remarks: Probably the holotype in the absence of other Morrison material. Poole (1989) stated the type is at MSU. Luperina (RWH 9393). Taeniocampa subterminata Smith, 1888a:476— 477. Type locality. “Northeastern and Middle States.” “It is much more common than alia, and I have found specimens in almost every collection I have seen.” Female, “N.Y. State” [M]; “Type”[C] [In excellent condition] Remarks: Todd (1982) designated the MSU specimen as the lectotype and stated that it “has been selected, labeled, and is now designated ...” I could find no lectotype label associated with the type. A lectotype label has been added to the specimen. A synonym of Orthosia revicta (Morr.) (RWH 10490). Heliothis suetus Grote, 1873b:117, pi. 3, fig. 10. Type locality. “Colorado Territory (coll. Theo. L. Mead, No. 7).” Male, "suetus Grt. Type. Colorado 298” [M] [In good condition, the antennae missing] Remarks: The holotype. In addition to containing another Grote name (see cali- fornicus), there were additional specimens included with the type but not from the type locality, since the description is based on only one specimen. The male marked with a Morrison label "suetus" is the only valid type in this collection (Hardwick, 1958, 1996). Hardwick (1958) recognized the male labeled in Morrison’s hand and marked "suetus" as the only remaining type from the series. Schinia (RWH 11088). 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 99 Hadena sujfusca Morrison, 1875d:61-62. Type locality. “Mass., Conn., Colorado.” Female, “Type”[C]; “Massachusetts 216” [M] [Well marked specimen in poor condition; the abdomen missing] Remarks: Syntype. The description clearly shows that a number of specimens were used to describe this species. In the original description, Morrison refers to “a poor specimen. . .from Massachusetts, in the collection of the Boston Society of Natural History. . .[and] several specimens at Nahant, Mass.” The locations of the other material is not known. A synonym of Apamea alia (Gn.) (RWH 9351). Mamestra teligera Morrison, 1874g[1875]:215— 216. Type locality. “Waco, Texas.” Male, “Adir. N.Y.”[T]; “Tepper”[ty]; ""teligera, Morr Type. Texas Oct 13 241 ”[M] [In good condition, the right antenna missing] Female, “ 19/10” [?] [In poor condition, fragments of legs glued back on spec- imen, abdomen glued back on] Remarks: Syntypes. I can find no explanation for the two locality labels on the type; my only guess is that a label from another specimen fell off another specimen (perhaps the female) and was inadvertently added to the type. The male is likely the type, given all the trademark features of a Morrison label, however, Lynette Warshow (pers. comm. 1995) states that syntypes of teligera are at MCZ. Their specimens are not the best candidates for type, however, since data cited indicate simply “Tex” with no mention of the name teligera associated with the specimen. A lectotype of this species should be designated, but until the MCZ material can be studied care- fully, such action should not be taken. Lasionycta (RWH 10393). Oncocnemis tenuifascia Smith, 1888b: 18— 19. Type locality. “Colorado. A single female in Mr. Tepper’s Collection.” Female, “Co.”[T]; “Tepper”[ty]; “Type”[C] [In very good condition] Remarks: Todd (1982) recognized this specimen as the holotype (RWH 10080). Lithophane tepida Grote, 1873a:27. Type locality. “Female Examined. Mass., Mr. H. K. Morrison, No. 908.” Female, ""tepida Grote, Type Camb. Ms. 3/18/74 74” [M]; “Type”[C] [In ex- cellent condition, the left antenna missing] Remarks: The status of this type is not known. Smith (1892:229) states that “a type is in the British Museum; another in the Tepper collection.” Poole (1989) states that the type is in the BMNH (RWH 9909). Schinia tepperi Morrison, 1875d:68— 69. Type locality. “Texas, September 15.” “Anterior tibia absent in the single spec- imen of the species before us, which is otherwise in good condition.” Male, “15/9”[?]; “Texas” [?]; “Type”[C] [All legs missing except right meta leg, antennae missing, wings in good condition] Female, “15/9”[?]; “Texas” [T] [In good condition] Remarks: Syntypes. It is clear that Morrison had but a single specimen before him at the time of description, however, there were two specimens with identical data and it is not possible to distinguish which is the type. Plagiomimicus (Poole 1989); Polenta (RWH 9755). 100 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Agrotis tepperi Smith, 1888a:452— 453. Type locality. “Montana. The unique female type is in Mr. Tepper’s collection.” Female, “Montana” [T]; “Type”[C] [The type in very good condition, the right antenna and meso leg missing] Remarks: Todd (1982) recognized this specimen as the holotype. Euagrotis (RWH 10910). Oncocnemis terminalis Smith, 1888b: 19—20. Type locality. “Colorado. A single female in Mr. Tepper’s Collection.” Female, “Col”[T]; “Tepper”[tyJ; “Type”[C] [Left meso and meta leg missing, in very good condition] Remarks: Todd (1982) recognized this as the holotype (RWH 10085). Mamestra thecata Morrison, 1875d:59-60. Type locality. “Glen Valley, near Mt. Washington, N. H., and Plymouth, Mass. The former specimen was received from Mr. S. H. Scudder, the latter from Mr. Edward Burgess, and was taken July 23, 1867.” Female, ""thecata, Morr Type, Plymouth M[s], July 23, 1867, 118”[M] [In fair condition, left forewing with nick, right antenna missing] Remarks: Syntype. The location of the other material is not known. A synonym of Anhimella contrahens (Wlk.) (RWH 10530). Hadena tortilis Grote, 1880c:46. Type locality. “Washington Territory. Coll, of Mr. Tepper.” Female, “W. T.”[M]; “Type”[C] [Left wings glued to thorax, right pro and meso legs missing; antennae missing] Remarks: It is not known how many specimens were utilized in the original de- scription. In the absence of any other Tepper material of this species, the above specimen is assumed to be the holotype. Fishia (RWH 9968). Agrotis trifasciata Smith, 1888a:460. Type locality. “Mt. Hood. One female specimen from Mr. Teppers Collection.” Female, “Mt. Hood”[T]; “Tepper” [ty]; “Type”[C]; “Euxoa Slide MSU No. ll ’’[ty] [In very good condition] Remarks: Todd (1982) recognized the MSU specimen as the holotype. Euxoa (RWH 10717). Agrotis unica Smith, 1890:70. Type locality. “I have somewhere named a very distinctly marked specimen with- out any confusing shades unica.'' Extracted from a longer quote in Todd (1982). Male, “N. Car.”[T]; “11”; “Tepper”[ty]; ""Agrotis unicus Type Smith”[S] [In good condition] Remarks: The holotype. While not intended as a published name, the mention of the name is ruled a valid description (Todd 1982). A synonym of Peridroma saucia; (RWH 10915); number not assigned in Hodges (1983). Homoptera uniformis Morrison, 1875h:148. Type locality. “Georgia. Received from my friend, Mr. George W. Peck, of Brook- lyn.” 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 101 Female, “Type”[C] [In poor condition, extra pin hole verdigris, only the right pro and meso legs complete, and portions of the left fore and hindwing tom] Remarks: Status uncertain. Unusual for a Morrison type, there is no label, only the yellow Cook type label. Smith (1893) stated the type is in the Tepper collection. A synonym of Zale declarans (Wlk.) (RWH 8691). Agrotis unimacula Morrison, 1874b[ 1875]: 166. Type locality. “Atlantic States.” Female, “Buffalo, N.Y.”[ty] [The antennae and right forewing partly broken] Remarks: The specimen was found unlabeled, in the general collection. Presum- ably Morrison had specimens from several states at the time of description. It may be a type if no other material is found in other collections. Poole (1989) stated that the type is at MSU. A synonym of Graphiphora haruspica (Grt.) (RWH 10928). Taeniocampa vegeta Morrison, 1875c[1876]:432-433. Type locality. “Dallas, Texas (Boll). From the collection of H. K. Morrison.” Male, “Dallas Tex. Boll”[ty]; “Type”[C] [In excellent condition] Remarks: The holotype. A synonym of Cissusa spadix (Cram.) (RWH 8592). Copipanolis vernalis Morrison, 1874b[ 1875]: 133-134. Type locality. “Massachusetts. One specimen taken on April 8, by Mr. Roland Thaxter.” Female, “Mass”[M] [In very good condition] Remarks: Syntype. This specimen was found unlabeled in the general collection. The status of this type is in question. In addition to this specimen, there was another male specimen collected in Massachusetts by Thaxter which matches in every way the original description of another Eutolype described by Grote (see rolandi). Poole (1994) indicated the presence of another specimen of vernalis at the MCZ, but was uncertain as to its validity. The MCZ specimen appears to be the better candidate. A synonym of Psaphida rolandi (Grote) Poole (1989, 1994); (RWH 10014). Hadena vulgivaga Morrison, 1874b[ 1875]: 144— 145. Type locality. Nebraska; New York. The two specimens before me of species are variable. The one from Nebraska, received through the kindness of Mr. G. M. Dodge, I consider the typical form.” Male, “Tepper”[ty]; “Glencoe Ne[b] 91”; “Type”[M] [In excellent condition] Remarks: From the description it is clear that there were two specimens before him; it is not clear where the other New York specimen is. A synonym of Oligia fractilinea (Grt.) (RWH 9406). Telesilla vesca Morrison, 1875L103. Type locality. “Texas; Wisconsin.” Female, ''vesca, Morr. Type. Wisconsin 261 ”[M] [In very good condition] Female, no labels. Remarks: The location of the Texas specimen is not known; in the absence of a suitable candidate the Wisconsin specimen is the holotype. A synonym of Galgula partita Gn. (RWH 9688). Agrotis wilsoni Grote, 1873c: 135. Type locality. “California.” 102 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Male, “Cal”[T] [In fair condition, antennae and pro legs missing] Remarks: This specimen was found in the general collection and not labeled as a type. Hardwick (1970) concluded that a specimen in the BMNH was probably not a type. He believed the type to be lost; this may be the type. Euxoa (RWH 10867). ACKNOWLEDGMENTS This project would not have been possible without the generous support of Lyman Briggs School, and the Department of Entomology at Michigan State University. Dr. John Rawlins (CMNH), Dr. John Franclemont (Cornell University), the late Dr. Roland Fisher (MSU) and Dorothy Frye (Michigan State University Archives) provided valuable historical information and insight. Dr. Frederick Stehr, Jim Zablotny, George Balogh, Dr. Charles Covell, Dr. Kauri Mikkola, Bob Kriegel, and Mogens Nielsen, aided this study with insight and assistance in several aspects of this study. Dr. Frederick Stehr, Jim Zablotny and Richard Leschen reviewed part, or all of the manscript. LITERATURE CITED Anonymous. April, 1889. The Speculum (Michigan Agr. Col. News). Coveil, C. V. 1970. A revision of the North American species of the genus Scopula Lepidop- tera: Geometridae). Trans. Am. Entomol. Soc. 96:101-221. Edwards, H. 1881a. A new genus and some new forms of North American Zygaenidae. Papilio 1:80-81. Edwards, H. 1881b. New genera and species of the Family Aegeridae. Papilio. 1:179-208. Eichlin, T. D. and W. D. Duckworth. 1988. Sesioidea: Sesiidae in Dominick, R. B. et al.. The Moths of America North of Mexico, Fasc. 5.1. Englehardt, G. P. 1946. The North American clear-wing moths of the Family Aegeriidae. Bull. U.S.N.M. 190:1-222, 32 plates. Ferguson, D. C. 1972. in Dominick, R. B. et al.. The Moths of North America North of Mexico, Fasc. 20. 2B, Bombycoidea (in part). Ferguson, D. 1985. The Moths of American North of Mexico, Fasc. 18.1 Geometroidea: Ge- ometridae (in part). Grote, A. R. 1873a. On the Noctuidae of North America. 6th Ann. Rep. Peabody Acad. Sci. 21-38. Grote, A. R. 1873b. VII A Study of North American Noctuidae. Bull. Buff. Soc. Nat. Sci. 1: 95-128. Grote, A. R. 1873c. VIII Descriptions of Noctuidae principally from California. Bull. Buff. Soc. Nat. Sci. 1:129-155. Grote, A. R. 1874a. Preliminary Catalogue of the Noctuidae of California, Part II. Can. En- tomol. 6:215-216. Grote, A. R. 1874b. Notes on American Lepidoptera with descriptions of twenty-one new species. Bull. Buff. Soc. Nat. Hist. 2:145-163. Grote, A. R. 1874c. New Species of North American Noctuidae. Proc. Acad. Nat. Sci. Phil. 1874:197-214. Grote, A. R. 1875. Preliminary list of the Noctuidae of California. Part III. Can. Entomol. 7: 25-28. Grote, A. R. 1880a. North American moths. Can. Entomol. 12:242-244, 254-288. Grote, A. R. 1880b[1879]. On Lithophane and new Noctuidae. Bull. U. S. Geol. Surv. Terr. 5:201-208. Grote, A. R. 1880c. New species of moths. Bull. Brook. Ent. Soc. 3:45-48. Grote, A. R. 1881a. North American moths. Can. Entomol. 13:15-17. 1997 TYPES OF LEPIDOPTERA IN THE TEPPER COLLECTION 103 Grote, A. R. 1881b. North American moths, with a preliminary catalogue of the species of Hadena and Polia. Bull. U.S. Geol. and Geogr. Surv. Terr. 6:251-211 . Grote, A. R. 1881c. Descriptions of four new species of moths. Papilio 1:4-6. Hardwick, D. E 1958. Taxonomy, life history, and habits of the elliptoid-eyed species of Schinia (Lepidoptera: Noctuidae), with notes on the Heliothidinae. Can. Entomol. Sup- plement 6, 116 pp 194 figures. Hardwick, D. E 1970. The genus Euxoa (Lepidoptera: Noctuidae) in North America. I. Sub- genera Orosagrotis, Longivesica, Chorizagrotis, Pleonectopoda, and Crassivesica. Mem. Entomol. Soc. Can. 67:1-177. Hardwick, D. E 1996. A Monograph to the North American Heliothentinae. No publisher given. 281 pp, 25 plates. Hodges, R. 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New genera and new species of North American Noctuidae. Proc. U.S.N.M., 10:450-479. Smith, J. B. 1888b. New species of Oncocnemis. Ins. Life. 1:18-20. Smith, J. B. 1890. Contributions toward a monograph of the insects of the Lepidopterous family Noctuidae of temperate North America-Revision of the species of the genus Agro- tis. Bull. U.S.N.M. 38:1-237. Smith, J. B. 1893. Catalogue of the Lepidopterous superfamily Noctuidae found in Boreal America. Bull. U.S.N.M. 44:1-424. Smith, J. B. 1895. Contribution toward a Monograph of the insects of the Lepidopterous Family Noctuidae of Boreal North America. A revision of the deltoid moths. Bull. U.S.N.M. 48:1-129, 14 plates. Tepper, F. 1881. Descriptions of New Lepidoptera. Bull. Brook. Entomol. Soc. 4:1-2, 1 plate. Tepper, F. 1883. Description of new moths. Bull. Brook. Entomol. Soc. 5:65-67. Todd, E. L. 1982. The noctuid type material of John B. Smith (Lepidoptera). U. S. Dept. Agr. Tech. Bull. 1645: 228 pp. Received 7 February 1997; accepted 21 June 1997. ADDENDA Dr. Tim McCabe had the opportunity to study the types after receiving the galleys, and suggests that both Mamestra teligra and Glaea sericea are names presently misapplied to other species. J. New York Entomol. Soc. 105( 1-2); 105-1 12, 1997 COMPETITION AND COEXISTENCE OF ANTS IN A SMALL PATCH OF RAINFOREST CANOPY IN PERUVIAN AMAZONIA John E. Tobin' Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138-2902 U.S.A. Abstract.-A structurally complex patch of forest canopy in Manu National Park (Peruvian Amazonia), consisting of two trees and eleven associated vines, was sampled using insecticidal fog. Approximately 62,000 ants were collected and sorted to species. The total biomass (dry weight) of adult ants in the sample was close to 49 g. Dolichoderus bispinosus, the dominant species in the sample, made up 64.2% of the ant biomass and 69.0% of the individual ants. The four most abundant species {Dolichoderus bispinosus, Dolichoderus decollatus, Azteca sp. 1, and Paraponera clavata) together comprised nearly 95% of the biomass and individuals in the sample. In spite of the clear dominance of the sample by a few species, a total of 85 species in 29 genera were found, making this the most species-rich point sample of a canopy ant fauna ever documented. Sampled at the level of a small number of trees, the rainforest canopy ant fauna reveals a pattern of remarkable species richness accompanied by strikingly low equitability in the rank- abundance distribution, or ecological diversity. A small number of species overwhelmingly dominate the ant assemblage but fail to exclude other ant species. Competition appears to limit the number of dominant species that can coexist in small areas, but a large majority of species present do not compete with the dominants and exist to a greater or lesser extent independently of them. The low ecological diversity observed in the sample may be in part a function of the spatial scale of sampling, and increased sampling should lead to a change in this pattern. The extent to which the species abundance distribution would become more equitable cannot be determined at this time. Finally, the structural complexity of the canopy may promote high species richness by creating microhabitat-linked species associations that effectively function as separate, non-competing ant assemblages. Key words: Formicidae, ants, rainforests, ecology, Amazon, Peru, canopy, biological di- versity. Until recently, information on the biology of arboreal ant assemblages was derived from isolated observations and sporadic collections carried out on treefalls. With the development of canopy fogging (Erwin, 1989a), more precise information has been gathered about the relative importance of ants in canopy arthropod communities (Moran and Southwood, 1982; Erwin, 1983b; Adis et al, 1984; Majer et al, 1990; Stork, 1991; reviewed by Tobin, 1995) and about the taxonomic composition of canopy ant assemblages (Wilson, 1987; Harada and Adis, 1994; Tobin, ms.; Tobin and Cover, ms.). The next logical stage in the examination of canopy ant assemblages involves characterizing them in terms of the relative importance of their component species. At this stage, however, little work has addressed this aspect of arboreal ant biology (but see Majer, 1990; Stork, 1991). Mailing address; John E. Tobin, Clifford Chance, 1 New York Plaza, New York, NY 10004. 106 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) In this paper I characterize for the first time the entire ant fauna of a small patch of tropical rainforest canopy in terms of both its taxonomic composition and of the numbers and biomass of the component species. This effort grew out of an attempt to determine the numbers of arthropod specimens and species per major taxon in a small, discrete patch of tropical forest canopy surrounding the Pakitza research sta- tion in Manu National Park, Madre de Dios, Peru (Erwin, 1989a, 1989b). Erwin reported already on these collections, and summarized data on the relative abundance of the major arthropod groups based on about two-thirds of the total sample (Erwin, 1989b). After nearly 82,400 arthropods had been sorted, 69.6% of these were ants. The rest of the arthropods were primarily beetles (9.1%), psocopterans (4.0%), dip- terans (2.5%), collembolans (2.2%), and spiders (2.0%). A preliminary report on the ants in these collections, based on partial samples, was published already (Tobin, 1991). This paper represents a final report based on the entire ant fauna, and contains a complete species list including changes resulting from recent taxonomic revisions (e.g., Shattuck, 1992). Little has been published on the relative importance of the component species in arboreal ant assemblages, and there are few studies of the taxonomic composition of point samples that can be used for comparison with this study. Wilson (1987) found 43 ant species in 26 genera in a single tree in Tambopata, Peru. Harada and Adis (1994), working in Brazilian Amazonia, sampled a number of trees and found as many as 77 species in individual trees. Tobin (ms.) found between 22 and 44 species in point samples collected along a transect in central Panama. These figures appear to be larger than comparable ones for Asia (Stork, 1991) and Australia (Majer, 1990). On first impression, the extreme skew in the ranked species abundance in this sample appears to be extraordinary. One species comprises the majority of the num- bers and biomass in the sample. The steep distribution includes a very long tail of species represented by numbers and/or biomass that are trivially small. This stands in contrast to the shallower, more equitable distribution of other tropical canopy arthropods such as beetles (Morse et al., 1988). I will argue, however, that the un- usual distribution of ant species abundances may be an artifact of the spatial scale of the sample. METHODS A discrete patch of canopy, consisting of two adjacent trees and eleven associated vines, was selected for sampling. The trees were identified as Matisia cordata (Bom- bacaceae) and Hirtella triandra (Chrysobalanaceae). In selecting the sampling area, the aim was to maximize the diversity of arthropods in the resulting point sample; thus the size and structural complexity of the trees, and the amount of associated epiphytic growth, were the main criteria used. A total of 93.6 m^ (1008 square feet) of plastic sheeting was hung in the understory to collect the falling specimens, and the selected area of canopy was treated with insecticidal fog (Erwin, 1983a, 1983b, 1989a). The resulting specimens were preserved in 70% ethanol for subsequent sort- ing and processing. In the lab, the arthropods in the sample were sorted to order, with the exception of the family Formicidae, which was separated from the rest of the Hymenoptera 1997 CANOPY ANTS OF AMAZONIA 107 (Erwin, 1989a, 1989b). Following this first sort, I sorted the ants to species. Species determinations were made at the Museum of Comparative Zoology (M.C.Z.), Har- vard University, on the basis of comparisons with available specimens and of pub- lished and unpublished keys. Voucher specimens are deposited at the M.C.Z. Following the identifications, biomass and the numbers of individuals were deter- mined for each species. For species with less than approximately 500 workers, the number of individuals was determined by direct counts or estimated from subsam- ples. Biomass was estimated based on the number of workers and the average dry weight of specimens of each species or of other species of similar size and body type. For the most abundant species, biomass was determined directly by weighing all the specimens of each species; the number of specimens per species was estimated from the total biomass of each species and the average dry weight of workers of that species. Samples were dried at 60°C until weight was stable. RESULTS A total of 85 ant species belonging to 29 genera and 5 subfamilies were found in the samples (Table 1). One species, Dolichoderus bispinosus (formerly known as Mon- ads bispinosa; Shattuck, 1992), dominated the sample, comprising approximately 65% of the biomass and 69% of the individuals in the collection (Table 2). Only four species {Dolichoderus bispinosus, Dolichoderus decollatus, Azteca sp. 1 , and Paraponera cla- vata) contributed more than 1.0 g to total biomass (dry weight). Other important species were Anochetus sp. nov., Pachycondyla sp. 1, and Camponotus sp. 3. The contribution to biomass of the majority of species in the sample was trivial, amounting to no more than a few milligrams (Fig. 1). In addition, 1.43 g of assorted ant brood were found in the sample; these have not been included in Table 2. In terms of numbers, the dominant species were D. bispinosus (43,200 workers), Azteca sp. 1 (14,500), D. decollatus (1,620), and Azteca sp. 2 (1,560). All other species were represented by fewer than 1,000 individuals. Among these nondominant species are, in decreasing order of abundance, Crematogaster sp. 1, Solenopsis sp. 4, Tapinoma sp. 1, Solenopsis sp. 3, 2iX\d Anochetus sp. nov. Of these, only Anochetus sp. nov. made a contribution of more than 1% to total biomass. Fifty-one species, or nearly 60% of the total, were represented by ten or fewer individuals. DISCUSSION A large body of evidence documents the critical role of competition in determining ant community structure in a wide variety of habitats (Hdlldobler and Wilson, 1990 and references therein). Numerous studies have documented dominance hierarchies in which certain species predictably displace others from foraging and nesting sites (e.g., Vepsalainen and Pisarski, 1982; Savolainen and Vepsalainen, 1988). Research on the spatial distribution of ant territories in tropical canopies has invoked com- petition to explain the commonly observed pattern of territorial exclusion by dom- inant species, known as an ant mosaic (Room, 1971; Majer, 1972; Leston, 1978; reviewed by Jackson, 1984). Under this view, colonies of dominant species partition the canopy into a series of mutually exclusive territories, and each of these codom- inant species is positively or negatively associated with a series of nondominant 108 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Table 1. Ant species collected in trees nos. 12 and 13, Zone 02/18/08, Pakitza, Manu Na- tional Park, Madre de Dios, Peru. A total of 85 species in 29 genera were found. PONERINAE Anochetus sp. nov. Gnamptogenys acuta G. concinna Hypoponera sp. 1 H. sp. 2 Odontomachus haematodus Pachycondyla cavinodis P. crenata P. unidentata P. sp. 1 Paraponera clavata Formicinae Brachymyrmex sp. 1 B. sp. 2 Camponotus bideus C. bradleyi C. heathi C. lancifer C. latangulus C. novogranadensis C. sp. 1 C. sp. 2 C. sp. 3 C. sp. 4 C. sp. 5 C. sp. 6 C. sp. 7 C. sp. 8 C. sp. 9 C. sp. 10 Dendromyrmex fabricii Myrmelachista sp. 1 Paratrechina sp. 1 PSEUDOMYRMECINAE Pseudomyrmex browni P. dendroicus P. godmani P. laevifrons P. oculatus P. pupa P. simplex P. tenuis P. tenuissimus P. sp. 1 Dolichoderinae Azteca sp. 1 A. sp. 2 Dolichoderus attelaboides D. bidens D. bispinosus D. decollatus D. diversus D. lutosus Myrmicinae Allomerus octoarticulatus Crematogaster sp. 1 C. sp. 2 C. sp. 3 C. sp. 4 Cyphomyrmex sp. 1 Leptothorax anduzei L. sp. 1 Ochetomyrmex sp. 1 Pheidole sp. 1 P. sp. 2 P. sp. 3 P. sp. 4 P. sp. 5 P. sp. 6 P. sp. 7 P. sp. 8 P. sp. 9 P. sp. 10 P. sp. 1 1 Procryptocerus sp. 1 Rogeria sp. 1 Sericomyrmex sp. 1 Smithistruma sp. 1 Solenopsis sp. 1 S. sp. 2 S. sp. 3 S. sp. 4 S. sp. 5 Strumigenys subedentata Wasmannia auropunctata Zacryptocerus sp. 1 Z. sp. 2 Tapinoma sp. 1 T. sp. 2 1997 CANOPY ANTS OF AMAZONIA 109 Table 2. Biomass and numbers of dominant ant species. Biomass Number of Species (grams) % Total biomass individuals* % Total numbers Dolichoderus bispinosus 31.51 64.2 43,200 69.0 Dolichoderus decollatus 11.68 23.8 1,620 2.6 Azteca sp. 1 1.67 3.4 14,500 23.1 Paraponera clavata 1.39 2.9 20 n.c.^ All other species'’ 2.80 5.7 3,300 5.3 Total 49.05 100.0 62,640 100.0 ' The numbers in this column are estimates based on subsamples. ^ P. clavata made a negligible contribution to numbers. '’This includes 81 species represented in the collections by workers and/or queens. species. Thus dominant ants are seen as the major determinants of ant community structure. Most ant mosaic research, however, has been carried out in tropical agroecosys- tems (principally in cacao plantations), which lack the structural and ecological com- plexity of natural forest canopies. To what extent principles derived from ant mosaic theory might be generalizable to natural forest canopies has been a matter of some controversy. Leston (1978) working in a forest near Bahia, Brazil, showed that dom- inant canopy ant species hold mutually exclusive territories and form a three-dimen- sional mosaic. In natural forest canopies, then, dominants are present and competition may determine the spatial distribution of dominant ant species inter se, as it does in Species Rank Fig. 1. Ranked species: Abundance plot. 110 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) tropical plantations. However, the little available evidence from natural forests does not tell us to what extent dominants may be the organizing agents vis-a-vis the nondominant species, which comprise the majority of the species richness. The ev- idence I present here, as well as evidence from natural forest canopies in Panama (Tobin, ms.), strongly indicates that dominant ant species do not determine ant com- munity composition as a whole in undisturbed tropical rainforest canopies. That dominant species do not always determine the composition and structure of canopy ant assemblages is suggested by a striking feature of the data presented in this and earlier, comparable studies. A surprisingly high fraction of the total ant species richness at a site can be found in any point sample collected at that site. For example, Wilson (1987) reported finding 135 species in an extensive series of col- lections across four forest types in the Tambopata Reserved Zone, Peru. In one, particularly diverse point sample he found 46 species, or nearly 35% of the entire fauna; only 6 of those species were not found in other samples from the area. Tobin (ms.), working in Panama, found that as many as 39% of the ant species in an extensive series of collections were found in a single point sample. Also, numerous species were present in a large number of samples irrespective of the dominant species in the sample. Thus dominant ant species are not linked to a few co-occurring nondominant species, as in classical ant mosaics. The picture that emerges is one of a mosaic of mutually exclusive dominant species superimposed upon an assemblage of species that are found throughout the forest but only in small numbers and bio- mass. This assemblage of nondominants appears, at least to some extent, to exist irrespective of the presence or absence of the dominant species. The diverse assemblage of nondominant species raises important questions. Wilson (1987), upon documenting the coexistence of 43 species of ants in one point sample, asked how it was possible for so many species to occupy the same site. With nearly twice that number of species in the present study, the question seems all the more pressing. One possibihty is that many nondominants are not territorial, and are coexisting by means that do not involve aggressive mutual exclusion, thus allowing higher local diversities. This is almost certainly true in part. The spatial scale of canopy foggings, however, may be too coarse to resolve ant mosaics of small nondominant species, to the extent that they exist. Competition may still play a role in structuring assemblages of nondominant ants that is presently invisible, given our current sampling methods. The data in the present study reveal a local ant assemblage in which one species preempts over two-thirds of the biomass space, the amount of biomass that an as- semblage can sustain given the energy entering the system and the conversion effi- ciencies of its component species (Tobin, 1994) (Fig. 1). While puzzling, this degree of dominance is a natural consequence of the spatial scales of both the dominant ant mosaics and the sampling regime in the present study. As seen already, a typical canopy sample, covering an area of approximately 100 m-, may reveal 25-50% of the total ant fauna of the area but only one or a small number of dominants. A second fogging sample would expose a few more nondominants in the tail of the species abundance distribution, but may reveal an entirely different dominant species. As similar samples are pooled, the tail of the distribution might only be between two and four times as long as that of the original sample, but a number of other dominant species would be revealed. The effect of this pooling would be to fill out the left side of the distribution without greatly increasing the length of the tail, thus 1997 CANOPY ANTS OF AMAZONIA 111 reducing its skewness and making it comparable to other, more typical distributions. Hence the scale of individual fogging samples, though large in terms of the number of individual arthropods collected, is too small to reveal the true species abundance distribution of an area. The ultimate explanation for high local ant diversity may reside in the extreme struc- tural complexity of the canopy environment itself (Lawton, 1983, 1986; Morse et al, 1985; Gunnarsson, 1992; see also Terborgh, 1985). Abiotic factors, such as temperature and humidity, range widely from the forest floor to the canopy (Parker, 1995), and different levels of the canopy are associated with different epiphytes and their attendant arthropod faunas. Epiphytic cacti may be common in the same forest that is thick with ferns near the ground. From the point of view of sensitive plants and small arthropods, then, the range of conditions along this altitudinal gradient results not in a single, ho- mogeneous canopy habitat but in a series of more or less distinguishable microhabitats superimposed upon each other from the understory to the high canopy. As in the case of the marine intertidal, alpha diversity in a tropical forest canopy may be more a function of a gradient of physical conditions over a small distance which can be finely partitioned, than of any one set of conditions in particular. ACKNOWLEDGMENTS I am grateful to Terry Erwin for sharing with me the ants from his study in Manu. Stefan Cover was invaluable in sharing his taxonomic expertise. Phil Ward kindly identified the Pseu- domyrmex in the samples. This work was supported in part by an N.S.F. pre-doctoral fellowship and by a Harvard Merit Fellowship. I am grateful to J. M. Carpenter, G. Chavarria. S. P. Cover, E. A. Stacy, and E. O. Wilson for comments on the manuscript. LITERATURE CITED Adis, J., Y. D. Lubin and G. G. Montgomery. 1984. Arthropods from the canopy of inundated and terra firme forests near Manaus, Brazil, with critical considerations on the pyrethrum- fogging technique. Studies Neotropic. Fauna Environ. 19:223-236. Erwin, T. L. 1983a. Tropical forest canopies: the last biotic frontier. Bull. Entomol. Soc. Am. 29:14-19. Erwin, T. L. 1983b. Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by insecticidal fogging. In\ S. L. Sutton, T. C. Whitmore and A. C. Chadwick (eds.). Tropical rain forest: Ecology and management: 59-75. Blackwell Scientific Pub- lications, Oxford, England. Erwin, T. L. 1989a. Canopy arthropod biodiversity: A chronology of sampling techniques and results. Rev. Peruana Entomol. 32:71-77. Erwin, T. L. 1989b. Sorting tropical forest canopy samples. Ins. Coll. News 2:8. Gunnarsson, B. 1992. 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Species number, species abundance and body length relationships of arboreal beetles in Bornean lowland rain forest trees. Ecol. Entomol. 13:25-37. Parker, G. G. 1995. Structure and microclimate of forest canopies. In\ M. D. Lowman and N. M. Nadkarni (eds.), Eorest canopies: 73-106. Academic Press, San Diego. Room, PM. 1971. The relative distribution of ant species in Ghana’s cocoa farms. J. Anim. Ecol. 40:735-751. Savolainen, R. and K. Vepsalainen. 1988. A competition hierarchy among boreal ants: impact on resource partitioning and community structure. Oikos 51:135-155. Shattuck, S. O. 1992. Generic revision of the ant subfamily Dolichoderinae. Sociobiology 21: 1-181. Stork, N. E. 1991. The composition of the arthropod fauna of Bornean lowland rain forest trees. J. Tropic. Ecol. 7:161-180. Terborgh, J. 1985. The vertical component of plant species diversity in temperate and tropical forests. Am. Nat. 126:760-776. Tobin, J. E. 1991. A Neotropical rainforest canopy ant community: Some ecological consid- erations. In: C. R. Huxley and D. E Cutler (eds.). Ant-plant interactions: 536-538. Oxford University Press, Oxford. Tobin, J. E. 1994. Ants as primary consumers: Diet and abundance in the Formicidae. In: J. H. Hunt and C. A. Nalepa (eds.). Nourishment and evolution in insect societies: 279- 307. Westview Press, Boulder. Tobin, J. E. 1995. Ecology and diversity of tropical forest canopy ants. In: M. D. Lowman and N. M. Nadkarni (eds.), Eorest canopies: 129-147. Academic Press, San Diego. Vepsalainen, K. and B. Pisarski. 1982. Assembly of island ant communities. Ann. Zool. Fenn. 19:327-335. Wilson, E. O. 1987. The arboreal ant fauna of a Peruvian Amazon forest: a first assessment. Biotropica 19:245-251. Received 1 February 1997; accepted 18 April 1997. J. New York Entomol. Soc. 105( 1— 2): 1 13— 120, 1997 RADIOGRAPHIC STUDY OF THE RESPONSE OF JAPANESE BEETLE LARVAE (COLEOPTERA: SCARABAEIDAE) TO SOIL-INCORPORATED MYCELIAL PARTICLES OF METARHIZIUM ANISOPLIAE (DEUTEROMYCETES) Rebecca C. Fry’, Linda A. Fergusson-Kolmes^, Steven A. Kolmes\ and Michael G. VillanF ' Department of Biology, Hobart and William Smith Colleges, Geneva, NY 14456 2 Department of Entomology, Barton Labs, New York State Agricultural Experiment Station, Geneva, NY 14456 ^ Department of Biology, University of Portland, 5000 N. Willamette Blvd, Portland, OR 97203. Author to whom reprint requests should be addressed. Abstract. — Mycelial particles of the entomopathogenic fungus, Metarhizium anisopliae (Metsch.) Sorokin, were grown in 0%, 0.025%, 0.25%, 1%, and 5% tryptone and incorporated into soil to test for effects on the behavior of Japanese beetle larvae, Popillia japonica Newman. In radiograph chambers, Japanese beetle larvae avoided soil containing mycelial particles be- ginning by 48 or 96 hr, and continuing until 192 hr (the end of the study). Differences in Japanese beetle responses to mycelia grown in various levels of tryptone were observed after 48 hr. Japanese beetle larval response was first observed at 96 hr in treatments of mycelial particles grown in 0.25% and 5% tryptone compared to 48 hr for particles grown in 0%, 0.025%, and 1% tryptone. Results show that fungal nutrition can influence the avoidance of M. anisopliae by Japanese beetles. Key words: Metarhizium anisopliae, Popillia japonica, tryptone, mycelium. Interest in the fungus Metarhizium anisopliae (Metsch.) Sorokin as a biological control agent against Japanese beetles has increased recently as studies show that this fungus can cause high levels of mortality in the laboratory (Krueger et al., 1992; Krueger et al., 1991). Field applications of entomopathogenic fungi, however, are often less successful (Rath, 1992; McCoy, 1990). One explanation is that many confounding parameters influence host-pathogen interactions in the field. Effective use of M. anisopliae as a control agent will rely on a thorough investigation of complex interactions among fungus, soil environment, and insect population (Villani and Wright, 1990). Villani et al. (1994) showed that Japanese beetle grubs avoid soil that contains mycelial particles of M. anisopliae grown in 0.25% tryptone. There has been a lack of evidence concerning the effects of fungal culture method on insect response to this fungus. Therefore, our study investigated the behavioral responses of Japanese beetle grubs to mycelial particles of M. anisopliae grown in 0%, 0.025%, 0.25%, 1%, and 5% tryptone. materials and methods Third instar Japanese beetle grubs from Horseheads and Tuscarora, NY, collected during September 1994, were used in all studies. Grubs were placed in moist soil. 114 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) covered with sod and stored at 10 ± 1°C, 0 h light, and 90% humidity until used in the experiment. To ensure that acceptably healthy grubs were used, they were re- moved from cold storage approximately fifteen minutes before all bioassays, and placed in individual cups. If no movement was observed, grubs were replaced. The soil used in the bioassays was unsterilized and screened using a 2 mm- screen. The soil was classified as a loam with a particle size distribution of 46.2% sand, 39.0% silt, and 14.8% clay. The organic matter content was determined to be 1.60%, the pH 6 .96 and the loose bulk density 1.19 g/cm^. Initial soil moisture was measured between 8.2-12.4% for the study. The strain of M. anisopliae used (ARSEF 2547 Entomopathogenic Fungus Culture Collection, USDA-ARS Plant Protection Unit, Cornell University, Ithaca, NY 14853), was originally isolated from a European Chafer {Rhizotrogus majalis) grub collected near Syracuse, New York. Mycelial materials were prepared and stored as described by Krueger et al. (1992). The x-ray system used for radiographs was a Hewlett-Packard Faxitron (Model 43855B, Hewlett-Packard Company, Palo Alto, CA). Voltage was set at 50 kVp for 5 seconds. A Kodak Industrex Instant Processor (Model P-1, Kodak, Rochester, NY) was used to process the radiographs (Villani and Wright, 1988). Mycelial particles prepared using the method described in Krueger et al. (1992) were used to inoculate SDAY (Sabouraud’s Dextrose Agar supplemented with 1.0% yeast extract) plates. After approximately 14 days, conidia were harvested by flush- ing the plates using 2 ml of sterile, distilled water plus 0.02% Tween 80 (Aldrich, Milwaukee, WI) and scraping gently with a sterile glass slide. Conidia and water were removed using a sterile pipette, then suspended in 10 ml of water plus 0.02% Tween 80 in a sterile test tube. The soaked conidia were stored for 24 hours at 4 ± 1°C. A mean number of 1.52 X 10^ (S.E. = 0.14 X 10^) conidia/ml was used in this experiment. Twenty g of dextrose (Fisher, Fair Lawn, NJ), 10 g of yeast extract (Amberex, Juneau, WI), and 3.75 ml of antifoam (Dow Corning, Midland, MI) were dis- solved in 1,000 ml of deionized water. After mixing thoroughly, 100 ml portions were dispensed into ten, 250 ml Erlenmeyer flasks. The following quantities 0.000 g (0%), 0.025 g (0.025%), 0.250 g (0.25%), 1.000 g (1%), and 5.000 g (5%) of tryptone (Marcor, Hackensack, NJ) were each added to two Erlenmeyer flasks for a total of 10 flasks. The flasks were sterilized for 20 minutes in the autoclave. When the liquid media had cooled in the flasks, 0.0124 g of Penicillin G (Sigma, St. Louis, MO, 1609 units/mg), 0.025 g of Streptomycin Sulfate (Sig- ma, St. Louis, MO, 763 units/mg), and 0.200 ml of the conidial suspension were added. The flasks were then placed in an automatic rotary shaker at 150 r.p.m. for five days at 25 ± 2°C. After five days, each mycelial culture was harvested by vacuum filtering the growth medium through filter paper (Fisher, Pittsburgh, PA, P8 “coarse”), using a Buchner funnel (approx. 9 cm). The resulting mycelial mats were immediately separated from the filter paper using forceps, placed in petri dishes and dried at room temperature for 48 hours. Dried mycelial mats were placed in zip-lock freezer bags and crushed by hand until all pieces were <2 mm^. Mycelial particles were stored at 0 ± 1°C until use in the radiographic study. Sixty chambers (Fig. 1) were constructed using 120, 17 X 100 mm polystyrene 1997 RADIOGRAPHIC STUDY OF JAPANESE BEETLE LARVAE 115 Fig. 1. Radiograph chambers. Chambers were created with dimensions of 17 X 17 X 200 mm. For treatment chambers, one side of each chamber was filled with soil containing mycelial particles and the other side with clean soil. Japanese beetle larvae were placed in center holes and observed over 192 hrs. culture tubes (Fisher, Pittsburgh, PA)-two for each chamber. A plastic test tube cap (Bellco “Kap-uts” Vineland, NJ) was modified to make it open at both ends. The two culture tubes were held together using the cap to attach them. After the two tubes were firmly held in place by the plastic cap, a 6 mm hole for larval introduction was drilled into the center of entire assembly. Using a heated paper- clip, two additional air holes (approx. 1 mm) were made, one at each end of the chamber. The chambers were then numbered from 1-60. The chambers were held horizontally during the experiments. To standardize the orientation of the cham- bers, one end was arbitrarily assigned as “top.” A random number generator (Microsoft Excel, Version 5.0) was used to determine which end of the chamber was filled first (fungus or clean) since variations in soil compaction can affect grub movement (Villani, unpublished data). In addition, the order of the chambers on x-ray plates and the position of fungus (fungus “top” or “bottom”) were randomly assigned. Treatment groups with 10 chambers/treatment were: mycelial particles cultures in 1) 0% tryptone, 2) 0.025% tryptone, 3) 0.25% tryptone, 4) 1% tryptone, 5) 5% tryptone, and 6) the corresponding control — noninoculated soil. All chambers, regardless of treatment, were filled before final assembly with 40 g of soil (20 g/tube) with a pinch of grass seed placed at both ends of the chamber. For those chambers which contained fungus, 0.15 g of mycelial particles were mixed thoroughly with 20 g of soil for a rate of 7.5 mg mycelial particles/g of soil. Using a small plastic funnel, one tube of each treatment chamber was filled with the soil/ fungal mixture, the other with clean soil. In the control chambers, both tubes were 116 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) m 0%TRYPTONE □ 0.025 % TRYPTONE ^ 0.25 % TRYPTONE □ 1% TRYPTONE g 5% TRYPTONE Fig. 2. Mean number of Japanese beetle larvae avoiding mycelial particles grown in 0%, 0.025%, 0.25%, 1%, and 5% tryptone after 1 hr, 48 hrs, 96 hrs, 144 hrs, and 192 hrs. Com- parisons with an expected even distribution that proved to be significant at P = 0.05 are indicated with an asterix (*). filled with clean soil. After the tubes were filled, they were reassembled and placed on x-ray trays. The chambers were secured using tape and stored horizontally. The holes of the filled chambers were covered with masking tape until infestation with the grubs to prevent water loss. After 24 hours, a third instar Japanese beetle grub was placed head first in the center hole of each chamber. Once the grubs had crawled completely into the hole of the chambers, the holes were again covered with tape. The chambers were stored at 25 ± 1°C, 60% humidity, and 16:8 (light: dark) during the experiment. Radiographs were taken at 1, 48, 96, 144, and 192 hr after infestation. The location of each grub was marked on the radiograph and recorded as “in” or “out” of the side of the chamber with fungus. Three replicates were performed at different times during February, 1995. The experimental design was a randomized complete block with fixed effects using treat- ment replicate as the blocking factor (Milton, 1992). ANOVAs were carried out using Microsoft Excel, Version 5.0. The analysis compared the total number of Japanese beetle grubs out of the fungus side of the chambers to an expected even distribution (50-50) at each sampling date. A Tukey test was used to compare the mean number of larvae avoiding fungus among treatments (Keppel, 1991). 1997 RADIOGRAPHIC STUDY OF JAPANESE BEETLE LARVAE 117 Fig. 3. Radiograph of mycelium containing chambers and eontrol chambers one hour after grubs were placed in eenter holes. Mycelium incorporated soil was placed in one half of each treatment chamber and is designated by a line. Grub location is indicated by a dot. A coin was used to mark the top of the tray. Grubs can be seen moving away from mycelium in a few of the chambers. RESULTS The mean number of Japanese beetle larvae avoiding incorporated mycelial par- ticles over time are shown in Fig. 2. After 1 hour, avoidance of the fungus was not observed (F = 0.723; df = 5, 10; P = 0.619) (Fig. 3). After 48 hr, the number of Japanese beetle grubs avoiding the mycelial particles grown in 0%, 0.025%, and 1% tryptone was significant (F = 5.234; df = 5, 10; P = 0.013). At 96 hr, the number of Japanese beetle grubs out of the side of the chamber with fungus was significant for all treatments (F = 6.135; df = 5, 10; P = 0.007). Japanese beetle grubs contin- ued to avoid all treatments of fungus after 144 hr (F = 5.389; df = 5, 10; P = 0.012), as well at 192 hr (F - 5.534; df - 5, 10; P = 0.011) (Fig. 4). A significant number of Japanese beetle grubs that entered the fungus side of the chambers after one hr, had left the fungus side after 48 hrs (F = 31.013, df = 1, 24, p = 0.000). The number of grubs which switched to the fungus side after 48 hrs, having entered the clean sides after one hour, was not significant (F = 3.045, df = 1, 24, p = 0.094). The side of the chamber filled first had no effect on Japanese beetle grub position (F = 0.653; df = 6, 12; P = 0.689). The number of grubs at the “top” of the control chambers did not differ from an expected even distribution (F = 0.955; df - 6, 12; P = 0.493). 118 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Fig. 4. Radiograph of mycelium containing chambers and control chambers eight days after grubs were placed in center holes. Grubs can be seen moved away from mycelium incorporated soil in all but one treatment chamber. DISCUSSION After one hour, Japanese beetle grubs did not avoid mycelial particles of M. an- isopliae in soil. The number of Japanese beetle grubs which avoided incorporated mycelial particles was significant after 96 hr. These results are consistent with pre- vious studies of grub behavior in response to mycelium of M. anisopliae (Villani et al., 1994). Most grubs that entered the sides of the chambers with incorporated fungus at the first sample date had moved into clean soil by the second sample date. Those grubs that entered the sides of the chambers with clean soil at the first sample date remained in the clean soil at the second sample date. These data support the hypothesis that grub behavior is influenced by the presence of M. anisopliae. After 48 hr, there was a difference between the avoidance by the grubs among the treatments. Japanese beetle larvae response to particles grown in 0.25% and 5% tryptone was delayed when compared to particles grown in 0%, 0.025%, and 1% tryptone. It has been hypothesized that repellent factors (i.e. metabolites) released by growing mycelium in the soil may be responsible for fungal avoidance by grubs (Villani et al., 1994). The variations in grub responses to the mycelium grown in different levels of tryptone after 48 hours suggest that the nutritional history of the fungus influences the repellent factor in mycelium. Previous studies have shown that the nutritional culture history of M. anisopliae influences fungal growth (Barnes et al., 1975; Campbell et al., 1978; Dillon and 1997 RADIOGRAPHIC STUDY OF JAPANESE BEETLE LARVAE 119 Chamley, 1990). There has been a lack of information on the behavioral responses by insects to these fungi grown under different nutritional regimes. Although the hypothesis was not tested directly, our study suggests that chemical factors associated with growing mycelium may be influenced by the nutritional culture history of the fungus and that larval Japanese beetles are repelled by these factors. These results may help to explain some inconsistencies in results that occur when M. anisopliae is used for pest control in the field. AC KNO WLEDGMENTS We would like to thank Nancy H. Consolie, Luann M. Preston- Wilsey, Wendy Heusler, Paul Robbins, and Robert J. Jarecke for technical support and Ariel Diaz for suggestions on data analysis (New York State Agricultural Experiment Station). Thanks to Diana Consolie and John Sherman at Hobart and William Smith Colleges in Geneva, New York, for help with construc- tion of the radiograph chambers. LITERATURE CITED Barnes, G. L., D. J. Beothel, R. D. Eikenbary, J. T. Criswell and C. R. Gentry. 1975. Growth and sporulation of Metarhizium anisopliae and Beauveria bassiana on media containing various peptone sources. J. Invertebr. Pathol. 25:301-305. Campbell, R. K., T. M. Perring, G. L. Barnes, R. D. Eikenbary and C. R. Gentry. 1978. Growth and sporulation of Beauveria bassiana and Metarhizium anisopliae on media containing various amino acids. J. Invertebr. Pathol. 31:289-295. Dillon, R. J. and A. K. Charnley. 1990. Initiation of germination in conidia of entomopatho- genic fungus, Metarhizium anisopliae . Mycol. Res. 94(3):299-304. Keppel, G. 1991. Design and Analysis. Prentice Hall, NJ. pp. 173-175. Krueger, S. R., M. G. Villani, J. P Nyrop and D. W Roberts. 1991. Effect of soil environment on the efficacy of fungal pathogens against scarab grubs in laboratory bioassays. J. Biological Control. 1:203-209. Krueger, S. R., M. G. Villani, A. S. Martins and D. W Roberts. 1992. Efficacy of soil appli- cations of Metarhizium anisopliae (Metsch.) Sorokin conidia, and standard and lyophi- lized mycelial particles against scarab grubs. J. Invertebr. Pathol. 59:54-60. McCoy, C. W. 1990. Entomogenous Fungi as Microbial Pesticides, pages 115-138. In: “New Directions in Biological Control: Alternatives for Suppressing Agricultural Pests and Diseases” R. R. Baker and P. E. Dunn (eds.) Liss: New York. Milton, S. J. 1992. Statistical Methods in the Biological and Health Sciences. McGraw-Hill, New York. Rath, A. C. 1992. Metarhizium anisopliae for control of the Tasmanian Pasture Scarab Ado- ryphorus couloni. pages 217-228. In: “The Use of Pathogens in Scarab Pest Manage- ment.” T. A. Jackson and T. R. Glare (eds.) Intercept Books, Andover. Villani, M. G. and R. J. Wright. 1988. Use of radiography in behavioral studies of turfgrass- infesting scarab grub species (Coleoptera: Scarabaeidae). Bull. Entomol. Soc. Am. 34: 132-144. Villani, M. G. and R. J. Wright. 1990. Environmental influences on soil macroarthropod be- havior in agricultural systems. Ann. Rev. Entomol. 35:249-269. Villani, M. G., S. R. Krueger, P. C. Schroeder, F. Consolie, N. H. Consolie, N. H., L. M. Preston- Wilsey and D. W. Roberts. 1994. Soil application effects of Metarhizium anisopliae on 120 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) Japanese Beetle (Coleoptera; Scarabaeidae) behavior and survival in turfgrass micro- eosms. Entomol. Soc. Am. 23:502-513. Zimmermann, G. 1993. The entomopathogenie fungus Metarhizium anisopliae and its potential as a biocontrol agent. Pestic. Sci. 37:375-379. Received 23 August 1996; accepted 4 April 1997. BOOK REVIEWS J. New York Entomol. Soc. 105(1— 2): 121— 125, 1997 The Wild Silk Moths of North America. A natural history of the Saturniidae of the United States and Canada. — Paul M. Tuskes, James P. Tuttle, and Michael M. Collins. 1996. Cornell University Press, Ithaca, N.Y. Hardcover, 250 pp., 30 color plates, 19 black-and-white illustrations, 22 drawings, 38 maps and overlays, 5 tables, 81/2 X 11. Cloth ISBN 0-8014-3130-1 $75.00. The Saturniidae are simply the non plus ultra of big, dazzling moths; the charis- matic megafauna of the insect world. Often mistaken for butterflies by non-ento- mologists, they are the specimens that collectors trot out for their relatives at Thanks- giving as examples of moths that are not small and brown (with occasional shades of gray). Satumiid workers never have to fish for complements or argue the attraction of their group by appealing to our appreciation for their subtle beauty. And as the most glaringly obvious examples of showy lepidoptera that are not butterflies, sa- tumiids are at once the poster children for invertebrate conservation efforts and the standard bearers for moth biologists who wish a more widespread audience for their work. Perhaps most significantly, they provide the model for the “Lunar Moth” that carried away Dr. Doolittle at the end of Richard Fleisher’s 1967 classic (actually, it resembles a Polyphemus more than it does a Luna; but at least 20th Century Fox got the subfamily right). Tuskes et al.’s volume represents the first synthesis of the natural history of North American satumiids that combines taxonomic treatments with high quality color pho- tographic plates of both adults and larvae. Pending the publication of LeMaire’s eagerly awaited volume on the Hemileucinae, no single treatment or series of treatments has covered the North American fauna since Ferguson’s (1971-72) Moths of North America (MONA) fascicle. In no case have the larvae or the life history details of the North American fauna been so thoroughly treated. Collins and Weast’s (1961) volume was taxonomically restricted, treating only the Satumiinae and Automeris; and the works of Dyar (1902), Packard (1905, 1914), and Michener (1952) were more limited with respect to their coverage of natural history and/or the taxa and life stages figured. No work has attempted to figure as much of the range of North American satumiid larvae since Packard’s (1905, 1914) excellent renderings of satumiid larvae at various instars. The Wild Silk Moths of North America thus fills a gap in the layman-friendly but thorough treatments of North American Lepidoptera. In a word, the summary of decades of life history work in this volume is masterful, and the large format, coffee-table style will make the book attractive to laymen as well as conservation biologists and expert and professional lepidopterists. The Wild Silk Moths of North America is divided into two primary sections, plus an introduction, two appendices, a bibliography and two indices. Part One (“Behav- ior and Ecology”) comprises seven chapters, of which the first four (Life History Strategies, Parasitism, Diseases of Saturniidae, and Populations, Species, and Tax- onomy) are strictly biological in focus. Chapter One (Life History Strategies) is a broad but excellent primer on the holobiology of satumiids. Chapters two and three 122 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) (Parasitism, Diseases of the Satumiidae) comprise less that six full pages of text between them, prompting one to wonder why their subject matter was divided out into separate chapter headings. Chapter two, at least, is complemented by Appendix 1 (pp. 217-223) which summarizes referenced records of parasitization and hyper- parasitization of the North American satumiid fauna. Chapter Four’s discussion of population biology, speciation, systematics, and “the subspecies problem” is trite and largely misled, relying heavily on what appear to be the a priorist paradigms associated with long-outdated Mayrian evolutionary taxonomy. Chapter 5 provides a number of important collecting tips to the novice; likewise Chapter 6 is an excellent road map to the powers and pitfalls of rearing. Chapter 7, “Silk Moths and Human Culture”, briefly summarizes the economic roles of satumiids in silk production, food, native american art, and as crop pests and medical hazards. Part Two, Species Accounts, comprises the bulk of the volume’s text and all of the plates. All three of the satumiid subfamilies with representatives in North Amer- ican are treated; 85 taxa in total, of which all are figured as adults and 75 as larvae. The larval plates are among the selling points of this volume. The quality of the color plates is excellent, and will enable virtually anyone encountering these organ- isms in the field to identify them painlessly. As most biologists will recognize, showy colors and spectacular designs in nature rarely go unaccompanied by fascinating behavioral, ecological, and evolutionary features. The Satumiidae are no exception, and Tuskes et al. do an excellent job of asking intriguing questions and, to some extent, providing novel data towards answering them. As it has for North American birds and butterflies, the systematics of North Amer- ican satumiids appears to have reached a point where most of the genera are more or less stable and where taxonomic changes consist largely of subspecific transfers, elevations, or synonymies. Although it does not purport to be a phylogenetic revi- sionary work, and does not describe any new taxa, it effectively revises the North American satumiid fauna at the alpha level. Only one change is made at the generic level: Syssphinx is resynonymized under Sphingicampa; the reverse was the single generic change in Lemaire (1988) (Rindge, 1989). The other taxonomic changes in Tuskes et al. are as follows: ten subspecies are sunk to within the nominate form {Eacles imperialis nobilis, Anisota stigma fuscosa, Anisota virginiensis pellucida, Anisota virginiensis discolor, Dryocampa rubicunda alba, Coloradia pandora lind- seyi, Hemileuca chinatiensis conwayae, Hemileuca nuttalli uniformis, Automeris io lilith, and Antherea polyphemus olivacea); one species is synonymized {Hemileuca artemis with nevadensis); two are elevated {Antherea oculea) or re-elevated {Fades os lari) to species status; and three species are reduced {Agapema anona platensis) or re-reduced {Hemileuca hera magnifica, Agapema anona dyari) to subspecific rank. By comparison, Ferguson (1971—2) described two new species {Sphingicampa blan- chardi and Agapema solita); four new subspecies {Anisota stigma fuscosa, Anisota virginiensis discolor, Hemileuca eglanterina annulata, and Automeris iris hesselo- rum)\ created two new combinations {Hemileuca hualapai and Hemileuca chinatien- sis); synonymized one variety {Pseudohazis hera var. arizonensis with Hemileuca nuttalli nuttalli) and one subspecies {Pseudohazis washingtonensis with Hemileuca nuttalli nuttalli); elevated three subspecies or varieties {Facies imperialis oslari, Sphingicampa heiligbrodti hubbardi, Sphingicampa [=Adelocephala] hogei var. montana) to species status; elevated one variety to subspecific rank {Sphingicampa 1997 BOOK REVIEWS 123 [=Adelocephala] quadrilineata occlusa); sunk two species to subspecies {Coloradia pandora lindseyi, Coloradia pandora davisi); and transferred one subspecies (Hemi- leuca eglanterina uniformis to H. nuttalli uniformis). It will be observed that many of the taxonomic decisions of Ferguson (and others) are reversed in Tuskes et al.’s treatment. Such is the nature of the systematic en- deavor: to pursue progress over stability. But one must wonder whether there con- tinues to be undue focus at the infraspecific and infrasubspecific levels, given that none of the most recent volumes on the Satumiidae have made much of an attempt at phytogeny reconstruction. (The most rigorous attempt to address the phytogeny of the Satumiidae to date was that of a hymenopterist [Michener, 1952] during his tenure at the American Museum of Natural History, and was strongly contested by lepidopterists [Forbes, 1952].) As a phylogenetic systematist, I find this emphasis not uncoincidental with the authors’ adherence to the biological species concept, and I fear that the shift away from macroevolutionary taxonomic questions to species- level taxonomic questions attributable to Mayr has been slow to reverse itself in the lepidopterist community. Belatedly, the volume suffers, in my opinion, from a chronic problem in its portrayal of species delineation and speciation. The author’s application of hybridization studies to the dehmitation of species is foreshadowed in what can only be seen as a rather outdated discussion of the species problem (pp. 3-4), and in the somewhat weak attempt in Chapter 4 to integrate evolution within populations and speciation. Throughout the book the discussion of specific (and infraspecific) dehneation (their “evolutionary view of the species”) is couched in Mayrian terms of interbreeding, hybridization, and sub- specific process-related assumptions. The Hemileuca maia complex is referred to as a “superspecies” (p. Ill); following Tuskes and Collins (1981), Satumia mendocino and S. walterorum are referred to as “semispecies” (P 163). Automeris io lilith is synony- mized under nominate io on the grounds that it “does not have a distribution pattern consistent with the subspecies concept” (p. 152). In fact, such terminology does not serve to clarify, but rather to confuse the logical relationship between alpha systematics and phylogeny reconstmction. It serves as well to misdirect the focus of systematic research away from recovering an underlying pattern from within which to test hypoth- eses, and towards an assumption-laden, process-oriented approach from within which hypothesis testing is impossible. One of the primary purposes of systematics is to provide an independent framework for the examination of biological processes. In contrast, Tuskes et al. (p. 32) state: “A consideration of the process of speciation is important in order to understand the con- troversy over what constitutes a species and what taxonomic rank should be assigned to a given population”. And later in the same paragraph: “Any theory of speciation must explain the transition from one stable, harmonious system of interacting genes to another such system.” Throughout the volume, the authors conflate the discovery op- erations used by systematists to delineate species with the process of speciation itself. Interbreeding and hybridization are thus seen as a tool for taxonomy rather than a phenomenon to be examined following character-based phylogenetic reconstruction. Re- productive isolation is repeatedly referred to as a “test” of species status. This approach, with its obligate reliance on the ability to produce fertile offspring, effectively abandons character-based systematic inference; indeed, one of the authors refers to morphological attributes as “indirect” (Collins, 1997). But the practice of systematics depends ulti- 124 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) mately on character-based species diagnoses. Only character-based diagnoses can lead to hypotheses of phylogenetic schemes and classifications if they are to be based on synapomorphyies. Species may certainly be “lost” by hybridization, which obscures phylogenetic pattern, but it is not acceptable to lump historically distinct entities because of what may (or may not) happen in the future. The authors’ adherence to the biological species concept is troublesome for other reasons. Tuskes et al. do not address the requirement that a species criterion must be consistent and operational if it is to apply to all organisms, and that not only are re- quirements of interfertization impossible to use consistently (they are irrelevant for asex- ual taxa, for example), the ability to interbreed may be plesiomorphic as well. In fact, Tuskes et al.’s Appendix Two “Satumiid Hybrids” (pp. 224-5), includes a number intergeneric crosses. It must be recognized that hybridization experiments such as those conducted by the authors are at best one-sided tests. Failure to produce fertile offspring may well be sufficient evidence from which to conclude that two putative species are distinct, as do Tuskes et al. for Eacles oslari and Antheraea oculea. But the ability to interbreed — the author’s primary justification for re-synonymizing Hemileuca artemis with H. nevadensis — cannot be invoked as de facto “proof” of legitimately character- based identification of conspecificity. Nonetheless, the authors are rather conservative in their taxonomic changes, sinking many more taxa than they elevated. Almost every such synonymy is based on refutation of allegedly diagnostic characters, the exceptions being Hemileuca artemis and Hemileuca nuttalli uniformis (whose distribution, the authors state on p. 144, “is inconsistent with that of a subspecies”). Perhaps the authors of this volume should either have included a review of higher level satumiid relationships (to the extent possible) or not devoted as much space trying to reconcile their notions of species with character data. Distributions, flight times, calling times, and host plant records (as well as at least one first-time larval description; Hemileuca hualapai, p. 108) are among the most valu- able aspects of the volume’s content. The volume is noteworthy for describing rangewide variation in flight times and development. The text is chock-full of useful basic natural history information and is at times downright entertaining. I was delighted to learn that the etymology of the Buck moth derives from the belief that the moth larvae developed in the heads of deer and that the adults fly from the bucks’ nostrils (p. 111). Likewise the anecdote of the polyphemus caterpillars’ sharing similar optical spectra with har- vested plums and getting sorted mechanically into the pmne processing machinery (p. 177) was most amusing. With almost 550 references, the bibliography is extensive, drawing on reports and season summaries by amateurs as well as revisionary and other empirical work by professionals. It serves as the most complete guide to the satumiid literature of which I am aware. Tuskes, Tuttle, and Collins are to be congratulated for their synthesis of North Amer- ican satumiid life histories. This volume is an up-to-date compilation of available natural history data that serves admirably both as a field guide and as an introduction to students, natural historians, and other biologists with an interest in silk moth biology. While their philosophy of systematics leaves something to be desired, their obvious dedication and diligence with respect to addressing basic natural history questions has resulted in one of the best lepidopteran natural history texts I have ever seen, and one that will no doubt set the standard for North American moth life history books for decades to come. — 1997 BOOK REVIEWS 125 Paul Z Goldstein, Dept, of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024. LITERATURE CITED Collins, M. 1997. Beyond cladistics — understanding the nature of species. Abstract. The Lep- idopterists’ Society 50th Anniversary Meeting. New Haven. Collins, M. M. and R. D. Weast. 1961. Wild Silk Moths of the United States. Saturniinae, III + 138p., fig. phot. Collins Radio Company, Cedar Rapids, Iowa. Dyar, H. G. 1902. List of North American Lepidoptera. USNM Bull. 52. Washington, D.C. Ferguson, D. C. 1971-72. In Dominick, R. B. et al.. The Moths of America North of Mexico, fasc. 20.2, Bombycoidea (Saturniidae), 275 p. + pp. XV-XXI, 30 fig, 22 pi. col. h.-t. E. W. Classey Ltd. And R.B.D. Publications Inc., London. Forbes, W. T. M. 1952. Reviews. The Saturniidae of the Western Hemisphere. . . by Charles D. Michener. Lep News, 6, pp. 109-111). Lemaire, C. 1988. The Saturniidae of America. Ceratocampinae. 480 pp. 64 pis, 379 text figs. Mus. Nac. Costa Rica. Soft cover. $80.00. Michener, C. D. 1952. The Saturniidae (Lepidoptera) of the Western Hemisphere. Morphology, phylogeny, and classification (Bull. Am. Mus. Nat. Hist., 98(5), pp. 335-502, 420 fig., pi. phot. h.-t. 5, 1 diagr., 19 tabls.) Packard, A. S. 1905. Monograph of the Bombycine Moths of North America, including their transformations and origin of the larval markings and armature, part 2. Family Cerato- campidae, subfamily Ceratocampinae (Mem. Natl. Acad. Sci., Washington, 9, 270 p., 19 fig., 60 pi. part col. h.-t.). Packard, A. S. 1914. Monograph of the Bombycine Moths of North America, including their transformations and origin of the larval markings and armature, part 3. Families Cera- tocampidae (exclusive of the Ceratocampinae), Saturniidae, Hemileucidae, and Brah- meidae. Edited by T. D. Cockerel (Mem. Natl. Acad. Sci., Washington, 12(1), IX + 516 p., 34 fig., 113 pi. part. col. h.-). Memoirs of the National Academy of Sciences. Volume XII Part III. Washington Rindge, F. H. 1989. Review of Lemaire, C. 1988. The Saturniidae of America. Ceratocampinae. Mus. Nac. Costa Rica. J. Lepid. Soc. 43(2): 154. Tuskes, P. M. and M. M. Collins. 1981. Hybridization of Saturnia mendocino and S. walter- orum, and phylogenetic notes on Saturnia and Agapema (Saturniidae). J. Lepid. Soc. 35: 1-21. J. New York Entomol. Soc. 105(1-2): 125-127, 1997 Click beetles: Genera of the Australian Elateridae (Coleoptera). — Andrew A. Calder. 1996. 401 pp., 420 figs, including 90 excellent habitus drawings. About $100.00 U.S. This appears to be an excellent reference to the genera of Australian click beetles. The illustrations are excellent, especially the habitus ones. The treatment for each genus starts with the original citation, synonymy if any, and the type species. This is followed with a very detailed description that is the equivalent of just over two pages plus about three more pages of illustrations. Finally, there is a short section on geographical distribution (world and Australian), a list of the Australian species. 126 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(1-2) biology, and comments. The distribution of Australian species is usually given as one locality and state or else state or area of Australia. There are 420 illustrations, including the 90 habitus drawings (one or two per genus). The other illustrations are of the male genitalia, female genital tract, and excellent SEM photographs of other parts (tarsi, heads, pronota, etc.). The subfamilies are briefly described each with a list of included genera and arranged in phylogenetic order, but the genera under each are arranged in alphabetical order, which I find distracting; on the other hand, everyone knows the alphabet! In the key, it would have been very useful to have included a page reference for each genus. The 660 Australian species are placed in 70 genera (48 are endemic) and eight subfamilies. A brief description and a list of genera is given for each subfamily. Fifteen new genera and seven new species are described; there are 13 species that remain unplaced and thus excluded from the fauna. Three other species, originally described as from Australia, could not possibly have come from Australia and thus are dropped from the list. Also dropped from the list is Lacon modestus, which was described from Australia, but has never been found “amongst the tens of thousands of Australian elaterids examined.” On the other hand, the more I looked at the “fine print,” the more I wondered if anyone really paid any attention to the smaller details. In many cases, the wording in the key does not match those in the descriptions. For example, in couplet 19 [the word prothorax should have been used, not prono- tum], “antennae inserted into a U-shaped depression (Figs. 33, 42)” [Fig. 33 is dorsal view of habitus and Fig. 42 is the hind wing!]. This sentence is redundant to the first one except there Fig. 43 is that of the female genital tract. Furthermore, antennae is plural and therefore depression should also be plural or use the word each — this is a fairly common mistake throughout the book. “Tarsomeres 1-4 each . . . (Figs. 37, 44)” [respectively these figures show the claw plus the distal part of tarsomere 5 and the female genital tract]. Also in couplet 19, the phrase “hind coxal cavity distally closed (rear wall of cavity not visible)” stumped me at first. Had the author stated “not visible ventrally” or even better, as in the descriptions (pp. 36, 40, 45), “hind coxal plate with distal width same as greastest proximal width” it would have made sense. In the descrip- tions, it is stated, “. . . ; hind coxal cavity closed distally (rear wall of cavity not visible).” Actually the rear wall of the cavity is visible from an oblique view if the hind leg is pulled backwards. Couplet 23, for Diadysis, “anterior section of pronotostemal suture not grooved” vs. “anterior portion of polished band along inner margin of hypomeral border.” Are these characters the same? Many of the illustrations do not match the key characters or even the description. For example, for Glypheus, compare couplet 25, “tarsomere 1 shorter than tarso- meres 2-4 combined” with the description on p. 182 that notes, “. . . slightly longer . . .” or measure these tarsomeres in Fig. 193 (slightly shorter). Fig. 194 (as long as). If this is because I am measuring them from different angles, then the author should have explained this in the introduction (dorsal or lateral view, from base to extreme tip including lobe, if any). The enlarged illustrations of the lateral lobes of the male genitalia are good, but 1997 BOOK REVIEWS 127 they certainly do not like the overall illustrations (compare Figs. 313 with 314 or 367 with 368). Many descriptions state “Pro thorax . . . anterior angles not strongly produced, only covering half of eye (at most),” yet many illustrations show anterior angles not touching the eyes (see Figs. 79, 229, 265, 355, etc.). This “deception” could be the result of the fact that the head may be extended or protracted in relation to the pro thorax. Lacon is listed as a valid genus from Australia (see p. vi), yet on p. ix it is removed from the list. The information on Lacon on p. 97 should have been placed on p. 384 along with the other excluded species? Despite my comments above, I still think that this a very useful book and worth the price. To prove the value of this book, the first Australian specimen I tried to identify was lacking the elytra, antennae, and tarsi, nevertheless, by thumbing through the habitus illustrations I was able to get a generic name, which I then verified by looking at the collection. — Edward C. Becker, Agriculture Canada, Ot- tawa, ON, KIA OC6, Canada. f r '- A ¥MT'^ -''Waititeli' i ' ^ ■ «^'S^r|^j?:0S5r3r ,. 'fm'ii -t , ' r >'"•«*;-*'’■ "< ^••''- 1 'i^r' >» ;(-82^ifb-.v •■•";• : ;•■ .>«*■*’' •.••■••#j».V?' kUi^SljcaslV '.v;'r»i*fl«i ■■:.■.>« i.-‘ Jf. •? t^«>K;^‘'!;’..nJSSi'-^' V* ' ‘I^^:,7,.4V‘ ,tj4§ . «l'i J $iti'||^Kiit ■ i; ^■•^^‘^ ■. -A. •S^!f'»^,-8..^‘'?.; > V >)! • tlW«l ' ,,;ty s', o, ft '"aV* '*’'ikf >♦’'-* . • f '. ,^r Ji ■ U^h i'W 'i^m ilNv 'f ■' , .-li) ’ nim^ ^'i^T'.'!^'-.i' .:'hi'.fLi.'' . :y-sS'^n**':L ^'-.<. .,;rp •’ ’ • > '< : ' >v • - • j -i. A\f 'hci%4 '■■' -S ■: ;■ '■■ <' ••• >!’i ■ *'‘'VaiV" U i . '... ■. * > ■ -'l, -■■ ■V’T^L, . ••■■■s’»..,fe-' 'V-'JJ i- 4 jf^i H ■ K '• s. ,'i. V ?x '■ ) ; .' tv ,'V:i^ ' I iia *'* ' * ‘i fc,-. ay- «■ INSTRUCTIONS TO AUTHORS The Journal of the New York Entomological Society publishes original research resulting from the study of insects and related taxa. 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Journal of the New York Entomological Society VOLUME 105 WINTER-SPRING 1997 NOS. 1-2 CONTENTS An analysis of the genus Stenomacra Stal with description of four new species, and some taxonomic rearrangements (Hemiptera: Heteroptera; Largidae) Harry Brailovsky and Cristina Mayorga 1-14 A new subgenus and two new species of the genus Machilinus (Meinertellidae, Archaeognatha = Microcoryphia, “Apterygota”, Insecta) from Mexico Helmut Sturm 15-23 A review of the genus Limnometra Mayr in New Guinea, with the description of a very large new species (Heteroptera: Gerridae) Dan A. Polhemus and John T. Polhemus 24-39 A new species of leaf-mining Oulema from Panama (Coleoptera: Chrysomelidae; Criocerinae) Fredric V. Vend and Annette Aiello 40-44 Nectarinella xavantinensis, a new neotropical social wasp (Hymenoptera: Vespi- dae; Polistinae) Sidnei Mateiis and Fernando B. Noll 45-49 Description of immature stages of two species of Pseudolampsis (Coleoptera: Chrysomelidae) and the establishment of a new combination in the genus Sonia A. Casari and Catherine N. Duckett . 50-64 Type specimens of Lepidoptera in the Tepper collection at Michigan State Uni- versity John H. Wilterding 65-104 Competition and coexistence of ants in a small patch of rainforest canopy in Peruvian Amazonia John E. Tobin 105-112 Radiographi'e study of the response of Japanese Beetle larvae (Coleoptera: Scar- abaeidae) to soil-incorporated mycelial particles of Metarhizium anisopliae (DeuteromyceteS) Rebecca C. Fry, Linda A. Fergusson-Kolmes, Steven A. Kolmes and Michael G. Villani 113-120 Book Reviews The Wild Silk Moths of North America. A natural history of the Saturniidae of the United States and Canada Paul Z. Goldstein 121-125 Click Beetles: Genera of the Australian Elateridae (Coleoptera) Edward C. Becker 125-127 SUMMER-FALL 1997 Nos. 3—4 I _ '^1 vT Vol. 105 Journal, New York Entomological Society (ISSN 0028-7199) Devoted to Entomology in General JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Editor; James M. Carpenter, Department of Entomology, American Mu- seum of Natural History, Central Park West at 79th Street, New York, NY 10024 Book Review Editor: James S. Miller, Department of Entomology, Amer- ican Museum of Natural History, Central Park West at 79th Street, New York, NY 10024 Publications Committee; James K. Liebherr, Cornell University, Ithaca, Chairman; David L. Wagner, University of Connecticut, Storrs; Alfred G. Wheeler, Jr., Clemson University, Clemson. 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Postmaster: Send address changes to the Nev/ York Entomological Society, % American Museum of Natural History, Central Park West at 79th Street, New York, NY I0024-.SI92. Known office of publication: American Museum of Natural History, New York, NY 10024. Journal of the New York Entomological Society, total copies printed 700, paid circulation 602, mail subscription 602, free distribution by mail 19, total distribution 621, 79 copies left over each quarter. @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). J. New York Entomol. Soc. 105(3— 4); 129— 1 39, 1997 MORPHOLOGICAL CASTE DIFFERENC;^ NEOTROPICAL SWARM-FOUNDING POLISllNll>wGi^^ PARACHARTERGUS SMITHII (HYMENOPTERA^ VESPIDAE) SiDNEi Mateus, Fernando Barbosa Noll and Ronaldo Zucchi Departamento de Biologia, Faculdade de Filosofia Ciencias e Letras de Ribeirao Preto, Universidade de Sao Paulo, 14040-901 Ribeirao Preto (SP), Brazil Abstract. — As part of a series of papers aimed at studying caste in epiponine wasps, this paper deals with Parachartergus smithii. The whole female population of a mature nest (131 workers, 14 intermediates and 5 queens) was measured and dissected. Canonical discriminant analyses indicated slight morphological caste differentiation. Mahalanobis distances for queen- worker, queen-intermediate and intermediate-worker were 10.33, 10.86 and 0.90, respectively. However, ovaries were much more developed in queens than in workers and intermediates indicating clear physiological differences. In addition, castes were distinguished by color pat- terns in the frons, clypeus, mandibles, gena and wings. An brief comparison of caste differences in the Epiponini reveals both allometric and non-allometric patterns. Key words: Polistinae, Epiponini, Parachartergus smithii, caste differences, multivariate analyses. The worker caste in social Hymenoptera consists of unfertilized females, normally incapable of fertilization and with undeveloped ovaries. In ants and honey bees workers are distinguished from the queen by discontinuous morphological characters. However, where such clear-cut distinctions are absent difficulties in distinguishing castes arise. For example, in the polistine tribe Epiponini, the presence of interme- diate females (Richards and Richards, 1951), which are characterized by ovary de- velopment in the absence of insemination, seems to be widespread. They appear in Protopolybia acutiscustis (cited as P. pumila, Naumann, 1970), P. exigua exigua (Simoes, 1977; Noll et al., 1996), Pseudopolybia vespiceps (Shima et al., in prep.). According to Richards (1971) the significance of the intermediates is practically unknown (but see Gastreich et al., 1993). However, there are direct observations on their frequent egg laying and oophagy in Protopolybia acusticutis (cited as P. pum- ila) and P. exigua exigua by Naumann (1970) and Simoes (pers. comm.), respec- tively. In epiponines, queens tend to be somewhat larger than workers. In Protone ctarina silveirae the queens are larger than workers (Shima et al., 1996b), but the most conspicuous size and color differences among epiponine wasps were found in some Agelaia spp. {=Stelopolybia), such as A. flavipennis (Evans and West-Eberhard, 1970), A. areata (Jeanne and Fagen, 1974), and A. vicina (Sakagami et al., 1996). In Pseudopolybia vespiceps caste differences are slight and intermediates are present (Shima et al., in prep.). In Parachartegus fraternus (Richards, 1978) and Charter- gellus communis (Mateus et al., 1996) differences are very slight or undetectable externally: queens are not obviously different from workers and intermediates. How- 130 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) ever, in some groups, queens are significantly smaller than workers in some char- acters and larger in others (Richards, 1971, 1978; Shima et al., 1994). According to Jeanne et al. (1995), who studied Apoica pallens, this is considered as non-size- based allometry, probably due to a reprogrammation in growth parameters (Wheeler, 1991). Such a pattern was detected also in Epipona guerini (Hunt et al., 1996), Pseudopolybia difficilis (Jeanne, 1996), Apoica flavissima (Shima et al., 1994), and Polybia dimidiata (Shima et al., 1996a). As the seventh report on our on going series related to caste differences in the swarm-founding polistine wasps, this paper deals with Parachartergus smithii. Other than the information summarized in Richards (1978) and the description of the initial steps of nest establishment (Ito, 1996), this taxon is poorly known. MATERIAL AND METHODS A mature colony of Parachartegus smithii (de Saussure) was collected (Dec. 6, 1994) in Cajuru, Sao Paulo State, SE Brazil. In the collected population 150 females and 25 males were found. The adults were put in Dietrich’s fixative for 72 hr and thereafter kept in 70% ethanol. In order to detect caste differences the following fifteen external body parts (Fig. 1) were manually measured in each female: (1) head width (HW), (2) head length (HL), (3) maximum interorbital distance (IDx), (4) minimum inteorbital distance (IDm), (5) gena width (GW), (6) eye width (EW), (7) mesosomal height (MSH), (8) mesoscutellar length (MTL), (9) mesoscutellar width (MSW), (10) pronotum width (PW), (11) meta- somal height (MH), (12) basal width tergum I (TIBL), (13) basal length tergum II (T2BL), (14) maximum width tergum II (T2ML), (15) partial length of fore- wing (WL). In addition, color patterns and other morphological characteristics were examined for 5 queens, 20 workers and 14 intermediates. We examined ovarial conditions by using stereomicroscope. In order to analyse insemination the spermatheca was removed and put on a slide in a solution of glycerine and alcohol (70%) in the same proportion. The presence of sperm cells was assessed by using a microscope. Statistical analyses including canonical discriminant anal- yses (CDA: Rao, 1973) and generalized Mahalanobis distances (D-) were per- formed with the SAS Program Package for PC computers. Kruskal- Wallis one way analyses of variance was used to detect differences for each character. If a difference was found, Dunn’s method for multiple comparisons was applied using the program Sigma Stat for Windows version 1.0. Two secretory organs, i.e., van der Vecht’s organ (sixth gastral sternal gland) and Richards’ gland (fifth gastral sternal gland), were examined in 5 queens, 10 intermediates, and 20 workers. RESULTS Nest architecture and colony composition. The nest was found at five meters height attached to a wooden post. The adult population comprised 150 females and 25 males. Ovarian dissection and the insemination test revealed the females to be 5 queens, 14 intermediates and 131 workers. The envelope was light gray with many brown and dark gray striated spots, producing a camouflage for the nest (Fig. 2A). It was 25 cm in length and 15 cm wide. The largest of thirteen combs had 136 cells and the smallest 7 cells; each comb was attached to the substrate by a single central 1997 CASTE DIFFERENCES IN PARACHARTERGUS SMITH II 131 Fig. 1. Illustration of the measured characters in the females of Parachartergus smithii. HW: head width, HL; head length, IDx and IDm; maximum and minimum interorbital distances respectively, GW: gena width, EW; eye width, PW: pronotum width, MH: metasomal height, T,BL: basal width tergum 1, T2BL: basal length tergum II, T2ML: maximum width tergum II, WL: partial length of the forewing. peduncle. The wood surface inside the envelope was partially covered by the same material used to build the combs (Fig. 2B). The total number of cells was 848, divided into 86 empty (most of them located in the periphery of the combs), 180 cells with eggs, 129 with small larvae, 80 with medium larvae, 61 with large larvae and 312 cells with pupae. The number 132 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) Fig. 2. Nest of Parachartegus smithii built on a branch. (A) bottom view with some wasps near the entrance and (B) an abandoned nest showing attachment of the combs to the substrate and the substrate surface partially covered by the same material used in the envelope and combs photographs taken by S. Mateus. of meconia was counted for each cell in order to evaluate the number of gener- ations produced. There were found 184 cells with one meconium, 206 cells with two, 152 cells with three, 74 cells with four and 68 cells with five meconia. In this way, the total number of individuals produced was estimated at 1688. Ovary development and spermathecal contents. The ovariole number was always three in each ovary, and the following types of ovarian development were dis- tinguished (Fig. 3): Type A subdivided into A, (N = 43) with filamentous ova- rioles which had no visible oocytes (Fig. 3A,) and A2 (N = 31) with some very small oocytes (Fig. 3A2). Type B was subdivided as B, (N = 57) bearing some young oocytes (Fig. 3B,) and B2 (N = 14) with one or more mature oocytes in each ovarioles (Fig. 3B2). Type C: (N = 5) with well developed and very long ovarioles with at least one mature egg, which were contorted inside the gaster (Fig. 3C). Insemination was confirmed only in females with type C ovaries. Color pattern dijferences. The principal color differences were on the head, meta- soma and wings. They were: 1 — mandibles light brown in queens, while chestnut in workers, although brown at the apex for both (Fig. 4, E vs. F); 2 — the clypeus. 1997 CASTE DIFFERENCES IN PARACHARTERGUS SMITH 1 1 133 Fig. 3. Kinds of ovary development found among the females of Parachartergus smithii: Type A (A1 — with filamentous ovarioles which had no visible oocytes and A2 — with some very small oocytes) Type B (B1 — bearing some young oocyte and B2 — with one or more mature oocytes in each ovarioles). Type C; well developed ovary present in inseminated fe- males. Scale bar = 1 mm. 134 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-^) supra-antennal area and paraocular spots light yellow in queens while yellow in workers (Fig. 4, A vs. B); 3 — the supra-antenal lateral area and the lower margin of the clypeus white in queens (pigment is absent) while yellow in workers (Fig. 4, A vs. B); 4 — the lower genal area close to mandibles white in queens (pigment is absent) and the upper part light-yellow, while in workers these are yellow and light- brown (Fig. 4, C vs. D); 5 — bands on metasomal terga narrow and light-brown in queens, wider and dark-brown in workers; 6 — radial cell areas of wings light-brown to chestnut in queens, while in workers this area is dark-brown to light-brown (Fig. 4, G vs. H). External glands. Richards’ gland was present and van der Vecht’s gland was absent in all analysed females. In queens the glandular tissue filled a third of the upper margin of the fifth sternum and was most abundant at the lateral borders. In workers and intermediates the amount of glandular tissue was smaller than that found in queens. Queen-worker differences in relation to morphometry. Among the fifteen analyzed characters all of them did not differ significantly among queens, workers and inter- mediates (Table 1). Canonical discriminant analyses (CD A) based on the fifteen analyzed characters indicated a slight difference among queens, workers and inter- mediates. Queens were located between —2 and 6 for CAN, values and workers and intermediates between —2 and 2.3 The following equation was used in order to calculate CAN,: CAN, = -21.8(HW - 2.29) - 9.57(HL 1.82) + 2.90(IDx 1.93) + 6.16(IDm - 1.00) - 6.79(GW - 0.53) - 13.29(EW - 0.56) + 11.89(MSH - 2.48) + 11.48(MTL - 1.68) - 12.42(MSW - 2.34) + 21.07(PW - 1.85) - 1.43(MH - 2.26) - 3.75(T1BL - 2.32) - 0.10(T2BL - 2.78) - 4.47(T2ML - 2.86) + 0.28(WL - 2.88) For determining CAN, mesoscutellar length (MTL), width of mesoscutum (MSW), head width (HW) and especially pronotum width (PW) were the most important among the analyzed characters. No differences in CAN2 values were found for the three groups. The following equation was employed in order to calculate CAN2: CAN2 = +15.41(HW - 2.29) - 0.99(HL - 1.82) - 6.37(IDx - 1.93) - 15.78(IDm - 1.00) + 22.97(GW - 0.53) + 10.12(EW - 0.56) + 3.16(MSH - 2.48) - 3.06(MTL - 1.68) - 11.89(MSW - 2.34) + 7.38(PW - 1.85) + 8.36(MH - 2.26) - 1.84(T1BL - 2.32) - 5.84(T2BL - 2.78) - 0.22(T2ML - 2.86) + 6.18(WL - 2.88). Minimum interorbital distance (IDm), head width (HW), mesoscutellar width (MSW) and especially metasomal height (MH) were the most important for deter- mining CAN2. Mahalanobis distances between queen/worker, queen/intermediate and intermedi- ate/worker were 10.33, 10.86 and 0.90 respectively, showing that queens are much 1997 CASTE DIFFERENCES IN PARACHARTERGUS SMlTHll 135 Fig. 4. Queen/worker differences in color patterns on clypeus and supra-antennal area (A, B), gena (C, D) (scale bar = 1 mm), mandible (E, F) (scale bar = 0.1 mm) and wing (G, H) (scale bar = 1 mm) of Parachartergus smithii. Numbers indicate color patterns; 1 dark-brown, 2 light-brown, 3 yellow, 4 light-yellow, 5 white and 6 chestnut. statistically significant (P < 0.0001). 136 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) $ (U c O is ^ § "S tn N T3 n 00 (N 00 On NO (N (N m o (N m in m O d d d d d d d d d d d d d d I I I I I I I I I I I I (n O NO On On On 00 (N 1^ m in O 00 q in O) (N 00 'It q 't; 1^ d On d NO NO d ^ s § 8 c Q O d OJ b T3 B ^ C O 5 ^ Oh O i 1 s 1 ^ ^_r (u o+ 2 g S Is » I f - S! o 09 (U . 00 2 g g B a o 3 s2 T3 ^ 2 "o 2 g S X B 5 (U OJ ^ (U s -g §■= 1 s s n ^ 73 "S w) 2 3 0 o (U g ^ 2 o Hi 3 w 2 S' 2:' h 02 03 .2^ •' O) O 3 02 O .2 2 ^ ° 3 2 02 3 o g 8 1 O O 02 3 -. 1 I £ ^ ■5 ^ '5 (U (U 2 00 2 d '3 > 2 > 3 2 3 0 3 0 ^ S' ^ ,S ,3 00 2;^ 00 g B ^ B ^ 3 U (X C3 O 3 (U to o X 2 £ O a o S 3^ 2 7j 2 3 8 2 o o separate (Fig. 16) teeth moderately separate (Fig. 18) (Fig. 17) 13a Metafemoral process $ At basal Vi of posterior edge At basal % of posterior edge At middle of posterior edge (Fig. 25) (Fig. 26) (Fig. 27) 174 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) Table 2. Character states shared between Homoiosternus canorum and Plesiosternus. No. Character H. canorum and Plesiosternus H. beckeri lb Meso and metatarsi S Short and thickened Long and narrow 2b Metafemora S Enlarged (Figs. 8, 9) Narrow (Fig. 7) The character 2c (Table 3) has been largely used to separate the genera of this subtribe into two groups, one with distinct and complete or briefly broken pronotal basal bead {Homoiosternus and Plesiosternus) and another with pronotal basal bead absent or only slightly impressed near the posterior angles (Moron, 1983, 1987); discovery of a second species of Plesiosternus and additional specimens of P. setosus with absent or feebly marked pronotal basal bead led to redefine the character states for Plesiosternus (Moron and Howden, 1992). In our opinion, this character should be secondarily used to separate Homoiosternus and Plesiosternus, due to its scarcely discernible character states. On other hand, similar shapes of male genitalia of the species of both genera not provide differences at the generic level (Figs. 19-24). We alternatively propose the characters Ic and 3c to recognize these genera (Table Figs. 1-9. Homoiosternus and Plesiosternus spp. 1-3. Male protarsus. 1) H. canorum\ 2) H. beckeri\ 3) P. setosus. 4-6. Male mesotarsal outer claw. 4) H. beckerv, 5) H. canorum; 6) P. setosus. 7-9. Male metatrochanter and femur (anterior to left). 7) H. beckeri (type); 8) H. canorum; 9) P. setosus. Scale = 1 mm. 1997 NEW MEXICAN HOMOIOSTERNUS 175 0+ 4-1 o . w 'H- DO £ o CO U £ 3 O 3 O D. ^ C-H 03 O 0.95 mm), longer scape (SI > 105). Sculpture with large pits with chagrination in between. First gastral segment ventro-anterior with a distinct collar which is bent ventrally. Head with a distinct carina along the posterior ventral face, which is darker than the adjacent surface. Ventral process of petiole in lateral view distinctly more darkly colored than the tergite boliviensis Mann - Small body size (TL < 0.95 mm), short scape (SI < 105). Sculpture fine and densely set. First gastral segment ventrally without a collar. Head with a bulge along the pos- terior ventral face, which, in lateral view is not darker than the adjacent surface. Ventral process of petiole in lateral view of the same color as the adjacent tergite 3 3. Head smaller (HL 0.57-0.60 mm, HW 0.35-0.36 mm), scape shorter (SL 0.35 mm) brujitae Agosti 1997 NEW PROBOLOMYRMEX FROM COSTA RICA 191 Figs. 1-4. Probolomyrmex guanacastensis n. sp. 1-2. Ergatoid female (paratype). 1. lateral view. 2. head in full frontal view. 3-4. Worker (holotype). 3. lateral view. 4. head in full frontal view. - Head larger (HL 0.63-0.65 mm, HW 0.39-0.40 mm), scape longer (SL 0.40-0.41 mm) guanacastensis n. sp. Probolomyrmex guanacastensis, new species Holotype: 1 worker, Costa Rica, Guanacaste, 8.5 km NW Bagaces, Plazuela Haci- enda Monteverde, 31 -VIII- 1996, S. O’Keefe, sift leaf debris. Holotype deposited at InBio, Costa Rica. Paratype: 1 queen, same data as holotype. Deposited in American Museum of Natural History. QUEEN. HL 0.63, HW 0.41, SL 0.40, TL 0.81, Cl 66, SI 97, Figures 1, 2 (1 examined). WORKER. HL 0.65, HW 0.39, SL 0.41, TL 0.79, Cl 0.60, SI 106, Figures 3, 4 (1 examined). Comments: This species is easily separated from P. petiolatus by the shape of the petiolar process, and from P. boliviensis by the smaller size, shorter scape, lack of a gastral collar, and similar coloration of the petiolar process and tergite. The ergatoid 192 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) female has eyes and ocelli, but flight sclerites are not developed. This species is phenotypically very similar to P. brujitae, but differs in having a relatively larger head and longer scape. Despite repeated sampling in July 1995, February 1996, and July 1996, only two specimens were collected of this genus from the type locality and none were col- lected in nearby localities of similar habitat. The specimens were found in sifted debris processed through a modified Berlese-Tulgren funnel from a riparian habitat that included Ochroma pyramidale, Albizzia samen, Anacardium excelsum, Ceiba pentadra, Hymenia courbaril, Bursera simiruba, Andira inermis, Spondias mombin, Sweetenia macrophylla, Cedrela odorata, Enterolobium cyclocarpum, Hirtella ra- cemosa, 2ind Ardesia revoluta. This site is unique in that numerous (>150) specimens of Homoconnus (Coleoptera: Scydmaenidae) were collected, while less than five were collected at all other sites sampled combined. P. guanacastensis is the first record of an ergatoid female in this genus, in which all the other known sexuals are of the normal winged type (Taylor, 1965). This species belongs to the exceptions among ants with ergatoid queens, which normally have large colonies (Peeters, 1997). The only completely collected colony so far included only 20 workers (Taylor, 1965: 360). ACKNOWLEDGMENTS STO would like to thank Gary and Das Stewart for permission to collect on their property, Gordon Frankie and Amigos de Lomas Barbudal for logistic support and housing, and Gordon Frankie for identifications of the major tree species in the type locality. Jack Longino was commenting on the danger of describing singletons. This work was funded in part by a Cali- fornia Agriculture Experiment Station Grant to Felix Sperling. LITERATURE CITED Agosti, D. 1994. A revision of the South American species of the ant genus Probolomyrmex (Hymenoptera: Formicidae). J. New York Entomol. Soc. 102:429-434. C. R. F. Brandao, J. L. M. Diniz, D. Agosti and J. H. Delabie. 1998. Revision of the Neotropical ant subfamily Leptanilloidinae. Syst. Entomol. (in press). Peeters, C., 1997. Morphologically ‘pbrnhive’ ants: comparative review of social characters, and the importance of queen-worker dimorphism. In: J. C. Choe and B. J. Crespi (eds.). The Evolution of Social Behavior in Insects and Arachnids: 372-391. Cambridge Uni- versity Press. Taylor, R. 1965. A monographic revision of the rare tropicopolitan ant genus Probolomyrmex Mayr (Hymenoptera: Formicidae). Trans. R. Entomol. Soc. Lond. 117:345-365. Received 27 January 1998; accepted 1 February 1998. J. New York Entomol. Soc. 105(3— 4): 193— 198, 1997 NEW SPECIES AND A NEW NAME FOR ANTILLEAN BUPRESTIDAE (COLEOPTERA) Henry A. Hespenheide Department of Biology, University of California, Los Angeles, California 90024-1606 Abstract. — Two species of Buprestidae are added to the Antillean insect fauna. Agrilus ja- maicensis is described as the second member of that genus known from the island of Jamaica. Taphrocerus chalumeaui is reported from Guadeloupe, taken on the leaves of the palm Euterpe globosa. Both species are more closely related to Central American forms than to known Antillean species. Agrilus hispaniolae is proposed as a new name to replace A. klapperichi Hespenheide, a primary homonym of A. klapperichi Obenberger. Fisher’s (1925) revision of Antillean Buprestidae reports only one species from the island of Jamaica now considered to belong to the genus Agrilus (Hespenheide, 1974), A. albicollis (Waterhouse), originally described as the type of the genus Par- adomorphus. Examination of collections during study of the genus Agrilus (Hespen- heide, 1974, 1979) yielded two Jamaican specimens that did not agree with these or the other species known from the Antilles. These specimens represent an unnamed species that is described below. More recently, F. Chalumeau has sent specimens of a species of Taphrocerus from Guadeloupe that is also unknown previously and described here. Agrilus jamaicensis, new species (Figs. 1-2) Description: Holotype female: Elongate, strongly flattened above; black with dark metallic blue green reflections throughout except for black median stripe on prono- tum and small purplish spot on upper middle of front; small spot of golden setae at anterior angles of pronotum, narrow line of white setae along elytral suture for distal %; 12.5 mm long, 3.0 mm wide. Head with front depressed along midline and transversely above middle; surface moderately rugose on upper half, finely punctate on lower half, sparse white setae along midline below middle and above epistoma; antennae reaching posterior angles of pronotum when laid alongside, serrate from segment 5. Pronotum widest at basal Vs; marginal and submarginal carinae separated along anterior %, median lobe of anterior margin very shallow; basal margin angulate- emarginate at middle of each elytron; disk moderately convex with shallow trans- verse depressions along base and before middle, narrowly depressed on lateral mar- gins; prehumeral carinae prominent, arcuate, joining marginal carina beyond middle; surface transversely, finely but distinctly rugose, not visibly punctate. Elytral margins undulate, gradually converging to tips; tips narrowly produced, divergent, narrowly angulate, each with prominent tooth on exterior edge; disk nearly flat with strong, small depressions at base and rather strong depressions along suture; 194 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Eigs. 1-2. Agrilus jamaicensis Hespenheide, n. sp.; line indicates scale. Eigure 1, dorsal view; Eigure 2, lateral view. Shaded regions indicate areas of dense setae. surface polished imbricate-punctate on outer halves, finely transversely rugose on inner halves. Prosternum glabrous, posterior process bilobed, expanded behind procoxae; pros- temal lobe subtruncate. Metacoxae shallowly emarginate on posterior margin, pos- terolateral angle produced; lateral half of metacoxae and adjacent portions of meta- sternum and epimeron with dark orange setae, medial half and adjacent metastemum with pale yellow setae. Abdomen with suture between sterna 1 and 2 strongly in- dicated at sides; apex of lateral portion of sternum 1 broadly rounded and expanded. 1997 NEW ANTILLEAN BUPRESTIDAE 195 sterna 1 and 3-5 with dark orange setae; pale yellow setae in broad triangular patches at anterior margins of vertical portions of sterna 1-5, patches of sterna 1 and 2 joined. Tarsal claws with inner teeth broader, rather long and parallel. Male unknown. Holotype: “Jamaica,” without precise locality (CASC). Paratype: “Jamaica,” without precise locality (BMNH). Comments: This striking species is very similar to A. dentifer Waterhouse, described from Mexico without more precise locality. A specimen of A. dentifer in my collec- tion from Barro Colorado Island, Panama, differs from A. jamaicensis as follows: A. dentifer is black with an indistinct spot of sparse white setae along the suture of the elytra at the middle, and lacks the extensive areas of pale setae on the metastemum and sides of the abdomen; the prehumeral carina is only narrowly separated from the marginal carina and the disk of the pronotum is visibly punctate and less strongly rugose; the front is very finely punctate below and nearly smooth on the upper half; the lateral portion of abdominal sternum 1 is only slightly expanded at its apex and the dorsal portions of sterna 3—5 have white rather than orange setae. Although these two species are distinct, they are clearly related, and are part of a larger group of species that includes A. apicatus Waterhouse, A. lentulus Waterhouse, A. cibarius Fisher, A. uvarovi Obenberger and others. All share a convex or weakly depressed pronotum with prehumeral carinae that join the marginal carinae, elongate elytral apices with strong external teeth and a narrow sutural stripe of setae, and similar colorations and patterns of setae on the lateral and ventral portions of the body. Agrilus albicollis from Jamaica is a black species with some golden reflections, measures less than 10 mm in length, and has the dorsal surface of the pronotum almost entirely obscured by white setae. It is certainly not closely related to A. jamaicensis, which is not very similar to the other Antillean Agrilus: A. denticornis Chevrolat from Cuba is a much smaller species (<5 mm); A. dominicanus Thomson from Cuba and Hispaniola is similar in size, but is uniformly and inconspicuously setose, has a depression along the midline of the pronotum, and has rounded and strongly dentate elytral apices (Fisher, 1925). That this large, striking species has not been collected recently probably only reflects the general lack of insect collections from the Antilles. Recent collections from Hispaniola have yielded unusual genera and new species, including another Agrilus (Hespenheide, 1990, and below), and systematic collecting in Jamaica or elsewhere in the West Indies is likely to be equally profitable. Taphrocerus chalumeaui, new species (Fig. 3) Description: Holotype Female: Moderately broad and flattened above, strongly shin- ing; olive green above, except darker spot on each elytron and scutellum and pro- notum with coppery reflections; front yellowish green; beneath more convex in cross- section, black with greenish reflections; length 4.3 mm, width 1.5 mm. Head with narrow medial depression, stronger above middle; surface strongly shagreened, irregularly punctate except for transverse impunctate band across middle of front, punctures small and shallow; sparse area of inconspicuous setae above epistoma, epistomal pores small, separated by own diameter; eyes not prominent. 196 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) 3 Fig. 3. Taphrocerus chaliirneaui Hespenheide, n. sp., dorsal view. Line indicates scale. Shaded regions indicate areas of dense setae. Pronotum 2 times as wide as long; moderately convex, narrowly transversely depressed along anterior margin and more broadly and deeply so along posterior margin but interrupted at middle, and irregularly so parallel to sides; short linear prehumeral protuberance in posterior angles; sides strongly angulate, widest behind middle; surface strongly shagreened, with ocellate punctures along all margins, spars- er on anterior margin and densest in posterior angles, each puncture with semire- cumbent seta. Scutellum strongly shagreened. Elytra at base wider than pronotum, widest at humeri, constricted behind humeri then shallowly arcuate to tips which are broadly, separately rounded and minutely denticulate; surface shagreened, more strongly so at tips and nearly smooth in oval area behind middle; punctures coarser at base becoming indistinct beyond the middle, second and fourth intervals slightly raised on basal half, suture and sixth interval more strongly raised for entire length (sixth interval subcarinate), especially toward apex, creating narrow triangular depression before apex; transversely depressed at 1997 NEW ANTILLEAN BUPRESTIDAE 197 base; setae condensed in spots at basal Va, at middle, and then along suture to tri- angular spot in subapical depression. Antennal grooves on prostemum rather broad and very shallow. Metacoxae con- spicuously setose. Abdomen beneath indistinctly shagreened, sparsely minutely punctate and setose; terminal segment broadly subtruncate at apex, apical groove following outline of apical margin, except slightly indented at middle, and overlain with row of long semi-erect setae. Holotype: Guadeloupe; Mome a Louis, 6.II.1972, F. Chalumeau, “adulte s[ur] feuille de Eut[erpe^ globosa” (IREC). Paratype: same data (IREC). This is the first species of Taphrocerus known from the Lesser Antilles and the first member of the genus from the Antilles definitely associated with a species of palm. Other Antillean Taphrocerus are associated with sedges (Cyperaceae), al- though only the Jamaican T. albomaculatus Fisher has been reared (T. W. Sherry and Hespenheide, unpublished). T. chalumeaui appears to be closely related to a group of species that in Central America includes T. albofasciatus Fisher (Panama) and several undescribed forms. This group is characterized by larger size and flattened shape, metallic red, blue or green colors instead of the black typical of most Taphrocerus-, laterally carinate elytra; and elytral setation patterns similar to that of T chalumeaui. T chalumeaui differs from the species mentioned above in having a color pattern that is olive- green rather than red or blue, in lacking definite lateral elytral carina, as well as in many finer details of morphology. The overall color and the pattern of setae on the elytra will separate it from all other known Antillean species. Agrilus hispaniolae, new name A. klapperichi Hespenheide, 1990:402 (primary homonym of A. klapperichi Oben- berger, 1940:183). Dr. Charles Bellamy has kindly pointed out that the name of my recently-described species from the Dominican Republic was preoccupied by that of a Chinese species not included in the most recent catalogue for the genus (Obenberger, 1936). ACKNOWLEDGMENTS The author is indebted to E Chalumeau of the Institut de Recherches Entomologiques de la Caraibe (IREC); Richard Thompson of The Natural History Museum, London (BMNH); David Kavanaugh of the California Academy of Sciences, San Francisco (CASC); John Kingsolver of the U.S. National Museum (USNM); and the Carnegie Museum, Pittsburgh, Pennsylvania (CMPI) for loaning specimens or providing assistance during visits. Dr. Charles Bellamy ad- vised me on nomenclatural matters. G. H. Nelson reviewed an early draft of the manuscript. Some financial support was provided in part by the UCLA Academic Senate. Margaret Ko- walczyk prepared the illustrations. LITERATURE CITED Fisher, W. S. 1925. A revision of the West Indian Coleoptera of the family Buprestidae. Proc. U.S. Natl. Mus. 65 (9): 1-207. 198 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Hespenheide, H. A. 1974. Nomenclatural notes on the Agrilinae (Coleoptera, Buprestidae): II. Agrilus. Entomol. News. 85:48-53. Hespenheide, H. A. 1979. Nomenclatural notes on the Agrilinae (Buprestidae): IV. Coleopt. Bull. 33:105-120. Hespenheide, H. A. 1990. New species of Buprestidae (Coleoptera) from the Dominican Re- public. Proc. Entomol. Soc. Washington 92:400-406. Obenberger, J. 1936. Buprestidae 5. In: W. Junk and S. Schenkling (eds.), Coleopterorum Ca- talogus 152:935-1246. W. Junk,’s Gravenhage. Obenberger, J. 1940. Ad regionis palaercticae Buprestidarum cognitionem addimenta. Sbornik Narodniho Musea v Praze, B (Prirodovedny) 2:111-189. Received 1 August 1996; accepted 26 March 1998. J. New York Entomol. Soc. 105(3— 4); 199— 205, 1997 THE ANT (HYMENOPTERA: FORMICIDAE) TYPES IN ARGENTINIAN COLLECTIONS Donat Agosti Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192 Abstract. — Type specimens of 114 species and subspecific ant taxa in Argentinian insect collections are listed, including 14 holotypes and 5 lectotypes. Most of the types are Neotrop- ical, with the exception of a series of Karavajev types from South East Asia. The two collection at the Division Entomologia, Museo Argentinao de Ciencias Naturales in Buenos Aires includes (MACN) and the Fundacion e Institute Miguel Lillo, Universidad Nacional de Tucuman in Tucuman (IMLA), house all ant types in Argentina. The collection of 50 Kusnezov types at IMLA, the Bruch collection and thirteen Karavajev types from SE-Asia in MACN is the most important holding because of the 14 holotypes and 2 lectotypes. The latter includes one holotype of each W. L. Brown, Jr., Th. Borgmeier, and W. Kempf. The Karavajev types are in a small separate box in MACN, which is labeled as follows: “Donacion Dr. Alejandro A. Oglobin” and “Hormigas tejedoras, Formici- dae, gen. Oecophylla, Polyrhachis, tipos del mirmecology Karawajew”. The box includes also 19 further species identified by Karawjew. Bruch’s collection is at MACN, but his extensive photographic documentation is at the Museo de la Plata in La Plata (MLPA). Type designation in myrmecology has in the past been rather loose. Ants, being social, led to a long upheld practice of not designing a holotype, but rather designate syntypes, types, including a series of specimens. The list is organized alphabetically following subfamilies and genera. Only the often fragmentary information provided on the labels are listed. The taxa are listed following the valid combination. If this does not agree with the label data, the orig- inal genus used on the label is provided in brackets. Types for which no correspond- ing description can be found are nevertheless listed. Further information on the species can be found in Bolton (1995). The abbreviations of the collections follows Arnett et al. (1993), which includes also the addresses of the collections. CERAPACHYINAE Acanthostichus femoralis Kusnezov Syntypes 17 workers, #10449; Argentina, For- mosa, Ing. Juarez, N. Kusnezov, viii. 1958, IMLA [manuscript name?]. Cerapachys silvestrii Mann Cotype 3 workers PR. Rio Pideras, Insular Expedition Station, F. Sein Jr., Feb. 5, 30. IMLA. In soil in sugar cane [manuscript name?]. DOLICHODERINAE Dolichoderus {Monads) andinus Kempf. Paratypes 2 workers, 1145, Peru Pichita, Caluga, 2,150 m, Weyrauch, 18.9.1960. IMLA. 200 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) Dolichoderus (Monads) lobicornis Kempf. Paratypes 3 workers. Blumenau, M. Wit- te, 10.12.1921. IMLA. Dorymyrmex (Dorymyrmex) emmaericaellus Kusnezov. Holotype and paratypes 6 workers. Bolivia, Cochabamba, La Paz. IMLA. Forelius (Neoforelius) tucumanus Kusnezov. Syntypes 2 females. #5617, Tucuman, N. Kusnezov. IMLA. ECITONINAE Edton mexicanum argentinum Borgmeier. Holotype 1 female. #32890, Roque Saenz Peria (Chaco), F. Ohnehider. MACN. Edton setigaster ^orgmdcY. Cotype 1 female. #11804, Bolivia. MACN. Neivamyrmex Edton hetschkoi oglobini Santschi. Typus 1 female. Argentina, Mi- siones Dr. A. A. Oglobin. MACN. Neivamyrmex hetschkoi Mayr. Cotype 1 worker. #2001, Argentina, Misiones, Est. Exp., Loreto, Dr. A. A. Oglobin. MACN. Neivamyrmex impudens W. M. Mann. Cotypes 3 workers. Honduras, Progresso, W. M. Mann. IMLA. Neivamyrmex laevigatum Borgmeier. Cotypes 2 workers, 1 male. #2580, Argentina, Rogne (?), S. Pena, N. Kusnezov, 13.6.48. IMLA. Neivamyrmex (Edton) pseudops grandipseudops Bruch. Typus 1 female. Argentina, Alta Gracia, La Granja, Sierras de Cordoba, C. Bruch. MACN. EORMICINAE Acropyga (Rhizomyrma) wheeled Mann. Cotypes 3 workers. Honduras, Lombardia. IMLA. Camponotus aguilerai Kusnezov. Syntypes 10 workers. Argentina, Jujuy, 6.1.1949. IMLA. Camponotus goeldii Forel. Syntype 1 worker. Brazil, Rio de Janeiro. IMLA. Camponotus guayapa Kusnezov. Holotype and paratypes 6 workers. La Rioja, Guay- apa, 12.8.1948. IMLA. Camponotus phytophilus Wheeler. Cotypes 4 workers. #2834/sk., Mexico, Guema- vaca, E. Skwarra, 6.26.1929. IMLA. Camponotus querdcola M. R. Smith. Paratypes 6 workers. USNM paratype #62025, Los Angeles, Tambark flat, T. C. Laurence, 7.15.1952. IMLA. Myrmelachista ina Kusnezov. Holotype and paratypes 36 workers. Argentina, Mi- siones, Manuel Belgrano, N. Kusnezov, 7.26.1949. IMLA. Myrmelachista schachovski Kusnezov. Syntypes 2 workers. Argentina, San Martin de Los Andes, S. Shajovskoi. IMLA. Myrmelachista ute Kusnezov Holotype and paratypes 20 workers. Argentina, Mi- siones, M. Belgrano, N. Kusnezov, 7.27.1947. IMLA. Myrmelachista vicina Kusnezov. Holotype (male), paratypes 1 male, 2 females, 1 worker. Neuqueu, Lago Cacar, Schachovskoi. IMLA. Camponotus mucronatus hirsutifrons [manuscript name?]. Types 6 workers. Guate- mala, Zacapa W. M. Wheeler, 12.13.1911. IMLA. Paratrechina dichroa Karavajev. Typus 1 worker. #2399, Prinseneiland, Karavajev. MACN (Karawajew). 1997 ANT TYPES IN ARGENTINA 201 Paratrechina koningsbergeri Karavajev. Typus 1 worker. #3869, Java, Jibodas, Ka- ravajev. MACN (Karawajew). Polyrhachis flavoflagellata Karavajev. Typus 1 worker. #2875, Java, Buitenzorg, Ka- ravajev. MACN (Karawajew). Polyrhachis keratifera Karavajev. Typus 1 worker. #2505, Amboina, Karavajev. MACN (Karawajew). Polyrhachis mystica Karavajev. Typus 1 worker. #2353, Java, Buitenzorg, Karavajev. MACN (Karawajew). Polyrhachis ochracea Karavajev. Typus 1 worker. #35090, Java, Buitenzorg, Kara- vajev. MACN (Karawajew). Polyrhachis rastellata javana Karavajev. Typus 1 worker. #2875, Java, Buitenzorg, Karavajev. MACN (Karawajew). Polyrhachis rotundiceps Karavajev. Typus 1 worker. #3004, Wammar, Ins. Am, Ka- ravajev. MACN (Karawajew). Polyrhachis schang gracilior Karavajev. Typus 1 worker. #2503, Amboina, Kara- vajev. MACN (Karawajew). Polyrhachis tibialis orientalis Karavajev. Typus 1 worker. #2439, Tifoc (Bocroc (?)), Karavajev. MACN (Karawajew). Polyrhachis tibialis robustior Karavajev. Typus 1 worker. #2583, Wammar, Ins. Am, Karavajev. MACN (Karawajew). Polyrhachis tubifex Karavajev. Typus 1 worker. #2359, Java, Buitenzorg, Karavajev. MACN (Karawajew). Pseudolasius carinatus Karavajev. Typus 1 worker. #2537, Aroe, Karavajev. MACN (Karawajew). MYRMICINAE Atta opaciceps Borgmeier. Paratype 1 worker. Brazil, Tapera, Pemambuca, F. Car- valho, 1.941. IMLA. Atta robusta Borgmeier. Paratypes 2 workers. Brazil, Rio de Janeiro, Sao Bento, F. Torres, 15.X.935. IMLA. Basiceros singularis F. Smith. Lectotype (designated by W. L. Brown) 1 worker. B. G., Kumpung, H. C. Lang. IMLA. Basiceros squamifer Borgmeier. Syntypes 3 workers. #5510, Rio de Janeiro, Angra dos Reis, Jussaral, Lopes and Lent. IMLA (ex coll. Borgmeier). Chelistruma lilloanus (Brown). Holotype and paratypes 3 workers. Argentina, Tuc- uman, N. Kusnezov, iii. 20.48. IMLA. Crematogaster hyperphyes Kusnezov. Paratype 1 female. #7080, Bolivia, Chapare, Marcus, 1.7.49. IMLA. Crematogaster hyperphyes Kusnezov. Paratypes 1 worker and 1 male. #7079, Bo- livia, Chapare, Marcus, 27.viii.49. IMLA [no type labels, but same blank red standard labels]. Lepthtorax pergandei Emery. Cotypes 2 workers. USNM Cotype #54078, USA, DC, Washington, 9.10. IMLA. Monomorium minimum cyaneum Wheeler and Mann. Paratypes 3 workers. #128, Mexico Hidalgo, Guerrero Mill, Mann. IMLA. 202 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Monomorium (Monomorium), minimum emersoni Gregg. Paratypes 3 worker USNM type #57292, USA, Texas, Austin, A. Emerson, iv.6.1937. IMLA. Neostruma zeteki Brown. Paratypes 2 workers. #5059, Panama, Barro Colorado Is- land, Zetek, Feb-March 43. IMLA. Pheidole eglobini Kusnezov. Paratypes 2 workers. #4699, Argentina, Misiones, Ya- cuy. IMLA [manuscript name?]. Cyphomyrmex hillbramus Kusnezov. Paratypes 2 workers. Argentina, Salta, R. So- ladillo, R.55-km 1442, 25.1.948. IMLA [manuscript name?]. Cyphomyrmex ulemli Kusnezov. Syntype 1 worker. #8849, Argentina, El Rey, Suita, 6.2.53. IMLA [manuscript name?]. Ochetomyrmex {Brownidris) argentinus (Kusnezov). Syntypes 1 female, 5 workers. Argentina, Misiones, Loreto, Dr. A. A. Oglobin. IMLA. Ochetomyrmex {Brownidris) argentinus (Kusnezov). Syntype 1 female. Tucuman, Trancas, ii.l947. IMLA. Ochetomyrmex {Brownidris) bolivianus (Kusnezov). Holotype 1 female. Bolivia, en- tre Wames y Montero, Sta. Cruz, Wygodzinsky, 27/28-1-1958. IMLA. Octostruma stenognatha Borgmeier and Kempf. Paratype 1 worker. Brazil, Nova Teutonia, 27°11'S, 52°23'W, 3-500 m, Fritz Plaumann, vi.l957. IMLA. Oligomyrmex {Erebomyrma) eidmanni (Menozzi). Syntypes 6 workers. Resistencia Chaco, 6.17.948. IMLA. Oligomyrmex {Erebomyrma) eidmanni (Menozzi) Syntypes 3 workers. Brazil, Men- des, H. Eidmann, 3.10.33. IMLA. Oligomyrmex stenopterus Kusnezov. Types 2 females. #6087, Argentina, Misiones, Est. Exp., Loreto, Dr. A. A. Oglobin. IMLA. Orectognathus clarki Brown. Paratype 1 worker. Australia, Victoria, Femtree Gulley, W. L. Brown, 1.1.58. IMLA. Oxyepoecus {Martia) inquilina (Kusnezov). Holotype 1 worker. #1832, Argentina, Tucuman, Jardin Ins. M. Lillo, N. Kusnezov, 2. v. 1948. IMLA. Oxyepoecus {Martia) minuta (Kusnezov). Holotype 1 worker. Tucuman, Quebrada Cainzo, N. Kusnezov, 8-iv.l948. IMLA. Pheidole bambusarum Forel. Cotypes 9 workers. Brazil, Sao Paulo, Grand bambou foret. Lutz. IMLA. Pheidole carapunco Kusnezov. Syntypes 1 soldier, 6 workers. #1295, Argentina, Tucuman, Carapunco, 2,600 m, N. Kusnezov. IMLA [1 worker lectotype desig- nated by E. O. Wilson]. Pheidole descolei Kusnezov. Lectotype and paralectotypes 9 workers. Argentina, Salta, Apolinario, Saravia, E E. Luna, 1948. IMLA [1 worker Lectotype designated by E. O. Wilson]. Pheidole gavrilovi Kusnezov Lectotype 1 worker, paralectotypes 1 female, 6 work- ers. #5137, Argentina, Misiones, Loreto, 19.vi.949. IMLA [1 worker Lectotype designated by E. O. Wilson]. Pheidole {Elasmopheidole) lilloi (Kusnezov). Holotype 1 female. Tucuman, 7.i.48, IMLA. Pheidole minutissima Kusnezov. Lectotype worker, paralectotypes 5 workers. #5348, Argentina, Misiones, Eldorado, N. Kusnezov. IMLA [1 worker Lectotype desig- nated by E. O. Wilson]. Pheidole {Elasmopheidole) subaberrans (Kusnezov). Paratypes 2 workers. #1679, 1997 ANT TYPES IN ARGENTINA 203 Salinas, 16.4.48; 1 female. #4586, Salta, Oran, 23.11.48; 2 females. Rio Calera, 25.4.48; 1 worker. #1792, ?. IMLA. Pheidole (Eriopheidole) symbiotica (Kusnezov) Paratypes 1 female, 1 male, 7177, La Picaga, N. Kusnezov, v.1951. IMLA. Pogonomyrmex anergismus Cole. Paratypes 2 females. #2911, USA, New Mexico, A. C. Cole, Sept. 12, 1961. IMLA. Pogonomyrmex guatemaltecus Wheeler. Cotypes 3 workers. MCZ-Cotype 20—24 20577, Guatemala, Zacapa, W. M. Wheeler, Dec. 12, 1911. IMLA. Pogonomyrmex longibarbis andinus Kusnezov. Paratypes 3 females, 2 males, 3 workers. #5712, San Antonio de los Cobres, 22.12.49. IMLA. Pogonomyrmex longibarbis andinus Kusnezov. Paratypes 3 workers. La Rioja, San Refugio del Penon, 4,000m, Monros, 1.47. IMLA. Pogonomyrmex marcusi lobatus Kusnezov. Paratypes (?) 3 workers. Bolivia, Co- chabamba, La Paz, N. Kusnezov. IMLA [unlabeled standard red type label]. Pogonomyrmex meridionalis leonis Kusnezov. Paratypes 4 workers. #5834, Santa Cruz, Canadon, N. Kusnezov. IMLA. Pogonomyrmex odoratus Kusnezov. Paratypes 3 males, 6 workers. #3553, Cerro Olto, 6.2.49. IMLA. Pogonomyrmex odoratus Kusnezov. Paratypes 1 female, 1 worker. Bariloche, 6.2.49. IMLA [unlabled red type label]. Pogonomyrmex rugosus Emery. Cotype 1 worker. USNM type #54071, USA, Cali- fornia, S. Jacinto, T. Pergand. IMLA. Procryptocerus marginatus Borgmeier. Paratype 1 worker. Brazil, Bahia, Uru^uca, P. Silva, 1947. IMLA. Procryptocerus montanus Kempf. Paratypes 12 workers. #1645, Brazil, SP, Campos do Jordao, Kempf, 14.xi.l956. IMLA. Solenopsis (Labauchena) acuminata (Borgmeier). Cotype 1 female. Luna. IMLA. Solenopsis basalis major Forel. Cotypes 6 workers. #12085, Argentina, La Plata, Bruch, 9.10.1912. IMLA. Solenopsis clyptemnestra bruchi Forel. Cotypes ? #12093/92, Argentina, La Plata, Bruch, 1892. IMLA. Solenopsis clyptemnestra orestes Forel. Cotypes 22 workers. Brazil, Ceara, Roche. IMLA. Solenopsis (Synsolenopsis) egregia (Kusnezov). Paratypes 4 females, 1 1 workers. #1298 Argentina, Tucuman, Tafi del Valle, N.K., 3.10.48. IMLA. Solenopsis {Synsolenopsis) eximia (Kusnezov). Syntype 1 worker. #2411, Uriburu, Formosa, 6.21.48. IMLA. Solenopsis globularia Forel. Syntypes 2 workers. IMLA. Solenopsis hammari carhuensis Forel. Cotype 1 worker. Buenos Aires, Bruch. IMLA. Solenopsis latastei hojfmanni Forel. Syntypes 5 workers, 7 females. IMLA. Solenopsis (Bisolenopsis) sea (Kusnezov). Syntypes 4 males, 4 females. #752, Ar- gentina, Santa Fe, Villa Ana, H. Willink, 2.1946. IMLA. Solenopsis {Paranamyrma) solenopsidis (Kusnezov). Paratypes 2 females. #7122, La Picaga, E. Rios, N. Kusnezov. IMLA. Solenopsis {Paranamyrma) solenopsidis (Kusnezov). Paratype 1 female. #4996, Mi- siones, Iguazu, N. Kusnezov, 25.7.49. IMLA. 204 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Solenopsis spei Forel. Cotypes 6 workers. Esperanza, A. Forel, in nest of Atta ce- phalotes. IMLA. Strumigenys planeti Brown. Paratypes 2 workers. Bolivia, Beni, Huachi, W. M. Mann, 1921-22, MuIIford Exp. IMLA. Strumigenys praecava Brown. Paratypes 3 workers. MCZ-Paratype #29293, Panama, Barro Colorado Island, W. M. Wheeler, 6.27.24. IMLA. Tetramorium (Triglyphothrixl) desertorum (Forel). Syntypes 8 workers. IMLA. Wasmannia williamsoni Kusnezov. I worker. #6060, Gral, Pico. IMLA [unlabeled red type label] Zacryptocerus (Paracryptocerus) borgmeieri (Kempf). Holotype and paratypes 6 workers. #5330, Misiones, Iguazu, N. Kusnezov, 29.7.49. IMLA. Zacryptocerus {Paracryptocerus) borgmeieri (Kempf). Paratype I worker. Misiones, Iguazu, N. Kusnezov. IMLA. Zacryptocerus {Paracryptocerus) cojfeae (Kempf). Paratype 1 female, I worker. Co- lombia, Cundinamana, Tibacuy, 18.iii.952. IMLA. Zacryptocerus {Paracryptocerus) cojfeae (Kempf). Paratype 1 worker. Colombia, Cundinamana, 1500m, Bemel + Mendoza, 13.3.1952. IMLA. PONERINAE Ectatomma regis Kusnezov. Holotype 1 worker. Argentina, Salta, El Rey, Nogues N. Kusnezov, 14. ii. 1953; paratype 1 worker. #6185, Argentina, Tucuman, Ruta 9 km 1335; paratype 1 worker, Argentina Rio Sali, 40 km of Tucuman 10.1.48 IMLA. Metapone hewitti Wheeler. Paratype 1 worker. F946, Malaysia, Borneo, Kuching, J. Hewitt. IMLA. Metapone tillyardi Wheeler. Cotype 1 female. MCZ-Cotype 1-2 20787, Mt. Tam- bourine, A. M. Lea. IMLA. Pachycondyla {Neoponera) goldbachi goyana (Kusnezov) Holotype and paratype 2 workers. #4792, Argentina, Misiones, Iguazu, 7.24.1949. IMLA. Probolomyrmex brujitae Agosti. Holotype, female. Argentina, Jujuy Aguas Blancas- Yaculica (Argentinian-Bolivian frontier), 22°43'44"S 64°22'25"W, 460 m, 25 Oc- tober 1994, D. Agosti & J. M. Carpenter. PSEUDOMYRMECINAE Pseudomyrmex acanthobia cocae Santschi. Typus 6 females, 3 males, 7 workers. Argentina, Alta Grande, La Granja, Sierras de Cordoba, C. Bruch, 1.922. MACN. Pseudomyrmex arborissanctae symbioticus Forel. Cotypes 6 workers, #12010, Co- lombia, Dibulla. IMLA. Pseudomyrmex {Pseudomyrma) championii haytiana paulina (Forel). Cotypus 1 fe- male, 1 worker Argentina, Bruch, 989. MACN. Pseudomyrmex flavidula laevivertex lizeri Santschi. Cotype 1 worker. #1144, Bolivia. MACN. Pseudomyrmex {Pseudomyrma) gracilis argentina (Santschi). Typus 2 workers. #2026, Argentina, Misiones, Loreto (Est. Exp.), Dr. A. A. Oglobin. MACN. Pseudomyrmex {Pseudomyrma) gracilis glabriventris (Santschi). Typus 1 worker, cotypus 1 female. #1170, Bolivia. MACN. 1997 ANT TYPES IN ARGENTINA 205 Pseudomyrmex (Pseudomyrma) gracilis sericata (Santschi). Typus 4 workers. Par- aguay. MACN. Pseudomyrmex oglobini Santschi. Cotypes (?) 9 workers. #2088, Argentina, Mi- siones, Loreto, Dr. A. A. Oglobin; Nido ne las tamas de lapacho (Tecoma Ipe) de 35 mts de altura - Patoreo (Grande) IMLA. ACKNOWLEDGMENTS This work was supported by a grant from NSF to Platnick. Willems was very helpful in organizing the visit in Tucuman as well as discussing the remain of other ant collections in Argentina. This compilation is part of a survey of Argentina, supported by the Argentinian National Parks through the necessary collecting per- mits. LITERATURE CITED Arnett, R. H., Jr., G. A. Samuelsen, and G. M. Nishida. 1993. The Insect and Spider Collections of the World. Elora and Eauna Handbook 11:310pp. Bolton, B. 1995. A New General Catalogue of the Ants of the World. Harvard University Press, 504 pp. Received and accepted 12 December 1997. J. New York Entomol. Soc. 105(3-4):206-214, 1997 NOTES ON THE GENUS THASUS (HEMIPTERA: COREIDAE) Carl W. Schaefer and Richard J. Packauskas' Department of Ecology and Evolutionary Biology, University of Connecticut, U-43, Storrs, Connecticut 06269-3043 Abstract. — New distributions are given for Thasus neocalifomicus Brailovsky and Barrera, T. gigas (Klug), T. acutangulus (StM), T. luteolus Brailovsky and Barrera, and T. rutilus Brailovsky and Barrera. Measurements of several instars and a key to the last three instars of the first three species are given. These new country records are given: T. acutangulus, El Salvador; T. luteolus, Panama; T. rutilus, Bolivia. The distribution of T. neocalifomicus is discussed, as are differences among its populations and between them and T. gigas. We conclude, tentatively, that T. neocalifor- nicus from the United States-Mexican landmass more closely resembles the neocalifomicus-gigas common ancestor than do T. neocalifomicus populations from Baja California Sur. The genus Thasus (Coreinae; Nematopodini) was revised recently by Brailovsky et al. (1995b). Since the publication of that revision, some additional data have become available, which we present here. These data are new distributional records for five species, and an account of the immatures of three of them. We also discuss the distribution of Thasus neocalifomicus Brailovsky and Barrera and distributional differences in the relative length of its third and fourth antennal segments; this rel- ative length is a feature that sets this species apart from others in the genus. NEW DISTRIBUTIONAL RECORDS OF THASUS SPP. Thasus acutangulus (Stal) — El Salvador: [new country record]: Mt. San Sal- vador, 4,000-6,400 ft; 2 mi down from Cerro Verde summit Guatemala: Antigua; Acatenango Honduras: [no other data] Mexico: Chiapas, nr. Slope of Cerro Bola; Chiapas, Municipio de Angel Albino Corzo, mountain rain forest; Municipio de Las Rosas, 3 km SE Aguacatenango, 1,671 m. Thasus gigas (Klug) — Guatemala: [new country record]: Cunea, 6,000 ft; Chichicastenango, 6,000 ft. Mexico: San Luis Potosi, 31 km SE of S.L. Potosi, 1,500 m; 2 mi SE of Pedro Montoya. Guadala- jara [no further data]. Chiapas, 5 km SE of Ji- totol, 1,676 m; Chiopas [sic], Escuintla Note: In the San Luis Potosi Pedro Montoya spec- imens, the fourth antennal segment is pale red- brown, like that of T. neocalifomicus (see diag- nosis of the latter in Brailovsky, et al. [1995b]) ' Present address: Department of Biological Sciences, Fort Hays State University, Hays, Kan- sas 67601-4099. 1997 NOTES ON THASUS 207 Thasus neocalifornicus — Thasus rutilus Brailovsky and Barrera — Thasus luteolus Brailovsky and Barrera — Mexico: Sinaloa, Los Mochis; 40 mi s. Culiacan. Sonora, Alamos; Llano; Minas Nuevas; Agua Zar- ca; E. of Carbo United States: Texas, Lyford (Willacy Co.) Bolivia: [new country record]: Prov. Sara Panama: [new country record]: Boquete, Chiriqui, 800 ft. The new distributional records for Thasus gigas and T. rutilus do not extend the ranges of these species notably. The distribution of T. neocalifornicus is considered in more detail below. We have also a specimen of T. neocalifornicus labeled simply “California, Wickham.” We cannot find a Wickham in California, and believe the label to be an error. If, as we suggest (Brailovsky et al. 1995b), the Costa Rican record of T. acutan- gulus is based on a misidentification, the present El Salvador record extends the range of this species somewhat further south. The new Panama record for T. luteolus, hitherto known only from Costa Rica (Brailovsky et al. 1995b), also extends this species’ range to the south. NYMPHS OF THASUS Among the specimens assembled for the revision were a few third through fifth instars, including some of the closely related genus (Brailovsky et al. 1995b) Pa- chylis. These nymphs were all dried, on pins. Because of their shriveled condition and because color varies markedly in Thasus (and probably in Pachylis too), we do not describe these nymphs, but record some measurements (Table 1). We give a key to separate the two genera and the first three instars of the three Thasus species available. The egg and the nymphs of Thasus gigas have been described and mea- sured by Brailovsky et al. (1995a). KEY 1. Humeral angles obtuse, but with small spine (if spine removed, angle obtuse); posterior border of pronotum with shallow median depression (Fig. 1); hind-tibial dilation asym- metrical on each side of medial tibial ridge (as in adult); one small but distinct tooth subapically on hind tibia Pachylis - Humeral angles sharp, acuminate, sometimes with spine (if spine removed, angle re- mains acuminate); posterior border of pronotum straight (Fig. 2); hind-tibial dilations symmetrical on each side of medial tibial ridge (as in adult); two small but distinct teeth of equal size subapically on hind tibia {Thasus) 2 2. Pronotum almost entirely dark, sometimes with pale central area; humeral angle with small acuminate point T. gigas - Pronotum not almost entirely dark, either lighter or with pale margins; humeral angle with large acuminate point 3 3. Pronotum dark with white or yellow lateral and posterior margins; fourth antennal segment 1.5 times longer than third T. acutangulus - Pronotum yellow to red on disc, this sometimes spreading to margins; fourth antennal segment subequal to or slightly longer than third T. neocalifornicus 208 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Table 1 . Measurements (mm) of Thasus nymphs. Thasus gigas Third instar Fourth instar Fifth instar (N = 3) (N = 3) (N = 5) Total length 15.5 (13.8-17.1) 17.8 (17.3-18.7) 25.2 (21.1-28.3) Width at widest point (abdomen) 8.1 (7.5-8.4) 10.4 (10.3-10.5) 12.5 (10.9-14.0) Antennal segments:! 4.6 (4.4_4.9) 6.2 (6.0-6.5) 6.2 (5.9-6.8) 2 3.7 (3.6-3.7) 5.1 (4.5-6.5) 5.0 (4.5-5. 8) 3 3.6 (3.6) 5.0 (4.9-5. 1) 4.8 (4.6-5. 1) width of 3 2.4 (2.3-2.5) 3.0 (2.9-3.2) 2.5 (2. 1-2.9) 4 3.6 (3.6) 4.9 (4.6-5. 1) 5.2 (4.9-5.5) Hind tibia: length 8.5 (8.4-8.9) 12.4 (12.2-12.8) 12.2 (11.5-13.0) width 2.0 (2.0) 2.9 (2.7-3.0) 2.7 (2.3-2.9) Thasus neocalifornicus Second instar Fifth instar (N = 2) (N = 8) Total length 10.7 (10.0-11.4) 25.7 (22.6-28.6) Width at widest point (abdomen) 5.5 (5. 3-5.6) 11.6 (9.8-13.6) Antennal segments: 1 3.2 (3. 1-3.2) 6.1 (5. 1-6.8) 2 2.5 (2.5) 4.9 (4.6-5.5) 3 2.6 (2.6) 4.6 (3.5-5.2) width of 3 1.2 (1.2) 1.9 (1. 0-2.9) 4 2.5 (2.4-2.5) 4.6 (4.3-4.8) Hind tibia: length 5.2 (5.2) 11.4 (10.4-12.1) width 0.9 (0.9) 2.5 (2.2-2.8) Thasus acutangulus Second instar Fifth instar (N = 1) (N = 4) Total length 10.6 21.6 (17.2-27.4) Width at widest point (abdomen) 6.2 10.7 (9.2-12.6) Antennal segments: 1 — 6.2 (6.0-6.4) 2 — 4.9 (4.7-5. 1) 3 — 4.8 (4.7-4.9) width of 3 — 2.6 (2.4-2.6) 4 — 6.6 (6.2-6.9) Hind tibia: length 6.0 11.4 (11.4-11.7) width 0.8 2.8 (2.6-3. 1) THE DISTRIBUTION AND POPULATIONS OF THASUS NEOCALIFORNICUS Thasus neocalifornicus had long masqueraded as T. gigas or T. acutangulus, until its status as a separate species was recognized (Brailovsky et al. 1995b). T. gigas is the sister species of T. neocalifornicus, and the two together compose the sister clade 1997 NOTES ON THASUS 209 0.6mm \ 2 h Figs. 1-2. Pronotum of fifth instar. 1, Pachylis sp. h = humeral angle. 2, Thasus acutan- gulus. h = humeral angle. of T. acutangulus + carchinus; the differences, similarities, and intraspecific varia- tions of these species have been described (Brailovsky et al. 1995b). Most of the several hundred Thasus neocalifornicus we have seen (and others recorded in Brailovsky et al. [1995b]), are from Baja California Sur, and the United States. Oddly, until now no one has noted that the United States T. neocalifornicus is almost completely restricted to the Tucson, Arizona, area, in southern Arizona (Pima, Cochise, and Santa Cruz counties); no specimen among the several hundred seen from the U.S. (including a large collection from Texas A&M University) has been found from west of this area, and only two specimens from east of it. Despite earlier listings of T. neocalifornicus (sometimes as Pachylis gigas) from New Mexico (e.g., Uhler, 1876; and see Brailovsky et al. [1995b] for summary), we have found no specimen from this state, although it begins only 100 miles east of the Arizona populations. The two specimens we have seen from east of Arizona were collected on August 19, 1975, in Lyford (Willacy Co.), Texas, which is about 40 miles north of Browns- ville. These specimens are somewhat darker than many others, but they fall well within the considerable range of color variation in this species. Another Texas population of Thasus neocalifornicus may be in the making. Dr. Walker Jones (USD A, Weslaco, Texas) writes (September 1993 and, in more detail, March 1995) that in 1992 he accidentally released a few females at Weslaco and, to his surprise, found nymphs the following spring; these he collected when they be- came adults but, if others avoided capture, a population may develop on Prosopis glandulosa, which is readily available. (Note: in his account of these bugs’ biology, Jones [1993] referred to them as Thasus “neomexicanus, ” a lapsus calami.) Welasco is some 30 miles south of Lyford. The Mexican populations of Thasus neocalifornicus are concentrated at the south- ern tip of Baja California Sur, in Miraflores, Sierra Laguna, La Paz, Todos Santos, El Triunfo, Cabo San Lucas, San Bartolo, and San Domingo; these are all within 50 210 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) miles of Cabo San Lucas, at the tip of the peninsula. In addition, we have seen specimens from Mulege, halfway up the inner coast, and from Rosario, on the outer coast some 150 miles south of the U.S. border. All these Baja California specimens were collected over half a century (1919-1971) by different people, most of whom were Americans. It seems likely then that these collectors did not travel only to the tip of the peninsula, but collected down its length and back up. In other words, the abundance of these bugs at the tip of Baja California Sur does not reflect collector bias; these bugs occur here in greater numbers than they do elsewhere on the pen- insula. They occur here in much greater numbers also than elsewhere in Mexico. We have specimens — only a few in each case — from only eight other localities, Los Mochis and near Culiacan (Sinaloa); and Santa Ana (Brailovsky et al. 1995b), Alamos, Llano, Minas Nuevas, Agua Zarca, and near Carbo (Sonora). Los Mochis is on the coast of the Gulf of California, not far from the Baja-tip populations; and Culiacan is some 100 miles south, also on the coast, near the tip of Baja California Sur. Santa Ana and Llano are equally close to the Arizona populations; Carbo is about 50 miles south of Santa Ana and roughly the same distance from the Gulf of California; but Alamos is far removed from any other population of T. neocalifornicus; we cannot find the other two Sonora localities on our atlases, but the entire state of Sonora is no more than 200 miles from the Gulf of California, and is across the Gulf from Baja California. We have examined representatives of these populations closely, seeking both intra- and interspecific relationships. We looked particularly closely at characters that sep- arate the species, especially T. neocalifornicus from T. gigas. We wondered if per- haps mainland T. neocalifornicus might more closely resemble T. gigas (also main- land) than do peninsular (Baja California) T. neocalifornicus. Genitalia. — The degree of spermathecal coiling and the shape of the tooth on the second valvula are the same in both Baja California and Arizona T. neocalifornicus, and differ from those in T. gigas: less coiling and more deeply bifid, respectively, in T. neocalifornicus than in T. gigas. With respect to the inner medial projection of the ventral rim of the male’s genital capsule, the variation between T. neocalifornicus and T. gigas is as great as that between T. neocalifornicus from Sonora and Sinaloa, and as that between T. neocalifornicus from Arizona and Baja California; also within this range of variation are specimens of T. gigas from Oaxaca, San Luis Potosi, and Guadalajara. The genital capsule of the single Texas male resembles those from Sonora more than those from elsewhere. Thus these genitalic characters neither support nor refute the idea that T. neocal- ifornicus from mainland Mexico (and Arizona) more closely resemble T. gigas than do T. neocalifornicus from Baja California. Color of fourth antennal segment. — This antennal segment in Thasus neocaliforni- cus, although usually reddish, may be as dark brown as that of T. gigas. The latter occurs farther south in Mexico, and the fourth antennal segment of the adult is longer than its third, never subequal. Also, the dark brown of the T. neocalifornicus fourth segment often pales to reddish distally. It is interesting in this regard that the third and fourth antennal segments of nymphal T. gigas are subequal (Table 1). The proximal half of the T. neocalifornicus second antennal segment may be paler than the distal half, rather than the more usual concolorous. Indeed, this segment in 1997 NOTES ON THASUS 211 several specimens from the Huachuca Mountains of Arizona is as red as in T. acu- tangulus, with which these specimens may be confused. However, antennal segments three and four are subequal in these specimens, proving them to be T. neocalifor- nicus. Humeral angles. — In Thasus neocalifornicus the humeral angles are produced as a small spine (Figs. 3-5). The spines of its sister species, T. gigas, are more blunt (Fig. 6). In the sister clade {T. acutangulus + carchinus) of both these species, the humeral angles of T. carchinus are more produced than they are in T. neocalifor- nicus, and the angles of T. acutangulus are sharp but not produced (Brailovsky et al. 1995b). The humeral angles of the Arizona and Mexican-mainland (Sinaloa, Sonora) T. neocalifornicus are somewhat less acute and produced (Figs. 3, 4) than are the angles of the Baja California Sur populations (Fig. 5). The former bugs’ humeral angles therefore resemble those of T. gigas slightly more closely than do those of the latter. Thus of the characters so far compared, only the humeral angles suggest that some populations of T. neocalifornicus may be more closely related to T. gigas than are other populations. This is insufficient evidence to suggest that T. neocalifornicus and its sister species T. gigas arose from their common ancestor on the Mexican main- land, although certainly the distribution of the two species supports such a sugges- tion. Ratio of third and fourth antennal segments. — Thasus neocalifornicus is alone in the genus in having the third and fourth antennal segments equal or subequal; the fourth is longer than the third in the other species, as it also is in the sister genus, Pachylis (Brailovsky et al. 1995b). We measured these segments in specimens from several T. neocalifornicus and T. gigas populations to see what differences there might be among populations, and if populations of one species geographically closer to the other might also be closer in third: fourth antennal segment ratios. The third and fourth antennal segments of Thasus neocalifornicus are equal (ratio of 1.0) to subequal (lowest ratio 0.85, from the Santa Rita Mountains and the Texas specimens) (Table 2). In T. gigas, on the other hand, the third is consistently shorter than the fourth (highest ratio 0.80, from Cuernavaca). The two segments from Baja California specimens are more nearly equal than are those from other localities’ specimens; in some of these Baja California specimens, the third antennal segment is actually slightly longer than the fourth (Table 4). All these Baja California specimens came from the tip of the peninsula; unfortunately, the specimens from further north were not available for measuring. The ratio of the U.S. (0.90) and of the mainland Mexican (0.87) T. neocalifornicus are lower (third and fourth less equal) than those of the Baja California (0.95) (third and fourth more nearly equal). The U.S. and mainland T. neocalifornicus ratios thus approach those of T. gigas (0.77) more closely than do those of the Baja California specimens. Conclusions. — The scant evidence presented here suggests (but only suggests) that the Baja California populations are isolated and have diverged more from the neo- californicus-gigas common ancestor than have the T. neocalifornicus populations on the United States-Mexican landmass. These latter populations are somewhat similar to T. gigas in the shape of their humeral angles and in the relative lengths of their third and fourth antennal segments. 212 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) 6.5mm Figs. 3-6. Pronota of adult Thasus. 3, T. neocalifomicus (Mexico: Sinaloa: Los Mochis). 4, T. neocalifomicus (U.S.: Arizona: Tucumcan). 5, T. neocalifomicus (Mexico: Baja California Sur: Miraflores). 6, T. gigas (holotype) (“Mexico”). 1997 NOTES ON THASUS 213 Table 2. Ratios of third to neocalifornicus and T. gigas. fourth antennal segments of different populations of Thasus Thasus neocalifornicus Third : fourth Locality ratio N Mexico: Sonora 0.85 7 Sinaloa 0.90 6 Mexico: mainland 0.87 13 Arizona: Tucson 0.95 3 Baboquivari Canyon 0.93 1 Patagonia 0.87 12 Santa Rita Mountains 0.85 2 “Arizona” 0.91 1 Santa Catalina Mountains 0.94 2 Fairbank 0.93 1 Arizona: all 0.90 21 Texas: 0.85 1 Baja California Sur Triunfo 0.97 16 San Domingo 1.05 1 Cabo San Lucas 0.92 2 San Bartolo 0.90 3 Miraflores 1.02 10 Sierra Laguna 0.96 7 Baja California Sur: all 0.95 39 Thasus gigas Third : fourth Locality ratio N Jalisco 0.75 11 Mexico Ternascaltepa, Cuernavaca 0.82 3 Chiapas 0.73 2 Hidalgo 0.77 3 San Luis Potosi 0.78 8 Mexico: all 0.77 27 ACKNOWLEDGMENTS We are very grateful to S. I. Frommer (University of California, Riverside), E. R. Hoebeke (Cornell University), K. J. Ribardo (California Academy of Sciences), and J. C. Schaffner (Texas A&M University), from whose institutions the specimens studied here were borrowed. J. Caira and her students, of this University, helped immeasurably with localities in Baja California Sur. The senior author thanks especially R. T. Schuh, of the American Museum of Natural History, for his hospitality, conversation, and access to Thasus. LITERATURE CITED Brailovsky, H., C. Mayorga, G. O. Leon and E. Barrera 1995a. Estadio ninfales de los coreidos del Valle de Tehuacan, Mexico (Hemiptera-Heteroptera) II. Especies asociadades a hui- zacheras {Acacia spp.) y mezquiteras {Prosopis spp.): Mozena lunata, Pachylis hector. 214 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) Savius jurgiosus jurgiosus y Thasus gigas. An. Inst. Biol. Univ. Auton. Mexico (Zool.) 66:57-80. Brailovsky, H., C. W. Schaefer, E. Barrera and R. J. Packauskas 1995b (dated 1994). A revision of the genus Thasus (Hemiptera: Coreidae: Coreinae: Nematopodini). J. New York En- tomol. Soc. 102:318-343. Jones, W. A. 1993. New host and habitat associations for some Arizona Pentatomoidea and Coreidae. Southwest. Entomol. Suppl. 16:1-29. Uhler, P. R. 1876. List of the Hemiptera of the region west of the Mississippi River, including those collected during the Hayden explorations of 1873. Bull. U.S. Geol. & Geograph. Surv. Terr. (2nd ser.) 5:269-362. Received 7 February 1997; accepted 22 October 1997. J. New York Entomol. Soc. 105(3-4):2 15-220, 1997 THE REAL IDENTITY OF DISCOELIUS STRIGOSUS COSTARRICENSIS BERTONI (HYM.: VESPIDAE: EUMENINAE) Bolivar R. Garcete Barrett Seccion Invertebrados, Museo Nacional de Historia Natural del Paraguay, Sucursal 1 Campus U.N.A., 2169 CDP, Central XI, San Lorenzo, Paraguay Abstract. — The identity of Discoelius strigosus costarricensis Bertoni ’= Zethus strigosus Saussure revised status, is solved and Zethus smidtianus n. sp. is described based on the mis- identification of costarricensis by Bohart and Stange, 1965 and subsequent authors. In 1925 Bertoni described Discoelius strigosus costarricensis, based on material from San Jose, Costa Rica. Forty years later, Bohart and Stange (1965) elevated the name costarricensis to species, based on material they believed they determined correctly, despite that they did not examine the type, considered to be — with the rest of Bertoni collection — locked in the basement of a bank in Asuncion, probably the Banco de Fomento (Willink, 1982; Carlos Aguilar pers. comm.). In 1991, thanks to the negotiations of Dr. Blanca Barrios, the Institute Agronomico Nacional (IAN) gave as donation to the Museo Nacional de Historia Natural del Paraguay, Inventario Biologico Nacional (IBNP) an insect collection that belonged to A. W. Bertoni. In 1995, thanks to the encouragement given by James Carpenter, Abraham Willink and Arnold Menke, I started to work this collection, finding in it a large amount of vespid type material described by Bertoni as well as interesting additional material that included specimens determined with unpublished names. The collection said to be deposited in the Banco de Fomento is no longer there and seem to be referable to the one IAN donated to the IBNP. Another insect collection that belonged to Bertoni was deposited in Puerto Bertoni (Garcete Barrett, 1996), until very recently, when John Kochalka and I brought it to the IBNP to be fully curated due its poor state of conservation. The complete restoration of these collections still awaits a long time, but in the mean while the discovery of the type of Discoelius strigosus costarricensis and a specimen identified under an unpublished name give light to a problem that may not be overlooked. Zethus strigosus Saussure, 1875 Zethus strigosus Saussure, 1875. Smithsonian Misc. Coll. 254: 42. Lectotype male, Orizaba, Veracruz, Mexico (Geneve). Zethus fortistriolatus Cameron, 1907. Entomologist 40: 82. Holotype female, Nica- ragua (London). Zethus mimus Zavattari, 1912. Arch f. Naturgesch. 78 (Abt. A), Heft 4: 64. Holotype female, “Brasilien” (Berlin). Discoelius strigosus costarricensis Bertoni, 1925. Rev. Soc. Cient. Paraguay (2) 1: 75. Lectotype female, San Jose, Costa Rica (Bertoni collection, IBNP), here des- ignated [examined], revised status. 216 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-^) The description given by Bertoni (1925) for Discoelius strigosus costarricensis is as follows: '"Foem. Atra, obscure sericea; abdominis primo segmento nitido, apice flavo; alis nigris, apice albis. clypeo truncate, striato-punchato[5'/c/]; fronte striata. Se le acerca la var d de Saussure (Am. Sol. Wasps, p. 42). La forma tipica de Mejico es mas clara y adomada, con las alas sub hialinas. Se asemeja en el peciolo a los Zetamenes W. Bertoni y en el color imita a Mischocyttarus smithi, etc. Col. Bertoni, San Jose de Costa Rica (C. Amer.).” This is a very short description but fits perfectly with a single female in the Bertoni collection, and labeled “Costa Rica, S. Jose”, “3167.”, ''Discoelius strigosus Sss. var. e ined.”. I also labeled it with the reference number “B. 209” as part of the recovery work. It bears no type label, but judging from the external evidence (Rec. 72B of the International Code of Zoological Nomenclature) there is no doubt this is a type specimen, and I designate it as Lectotype according with Rec. 73F and Art. 74b of the International Code of Zoological Nomenclature, taking into account that Bertoni did not mention how many specimens he saw. The specimen fits with the key and description given by Bohart and Stange (1965) for Zethus strigosus Saussure, but is darker, with yellow marks as follows: conspic- uous mandibular band and subapical band on tergum I, this last suddenly becoming thinner and briefly disappearing medially; less conspicuous are: scrobal spots, tiny spot in the inner orbit, tiny spot behind the posterior angle of the eye, mostly beneath flagelomeres II-X, a pair of indistinguishable tiny humeral spots, a pair of thin longitudinal bands on the posterior face of the propodeum, and lateral, almost in- distinguishable, indications of subapical bands on terga II, IV-V and sterna III, IV- V, as well as basolateral spot on tergum VI. The wings are more or less black, mainly along the costal area of the forewing, this last having a whitish tip. I compared this specimen with reference material of Zethus strigosus determined by Bohart and Stange: Mexico: Nayarit, 6 mi. E. San Bias, 4. ii. 64, 1 female (E. I. Schlinger col.) y 1 male (M. E. Irwin col.), UCD; Veracruz, Jalapa, 1 female, UCD; Oaxaca, 12 mi S. Chivela, 18. viii. 1958, 1 male (L. A. Stange y A. S. Menke cols.), UCD; Panama: Potrerillos, 5. v. 35, 2 males UCD. I was unable to find any structural difference between the Lectotype of strigosus costarricensis and the reference material. About the color pattern I must say that it is still in the range of variation within the species, existing only a remarkable re- duction of the yellow markings, the male from Nayarit is the closest in this respect. The only really remarkable difference is the wing pattern, almost uniformly hyaline yellowish in all the additional specimens. Zethus smidtianus, n. sp. [Zethus costarricensis: Bohart and Stange, 1965. Univ. Calif. FTib. Entom. 40: 97. Subsequently: Stange, 1978. Acta Zool. Lilloana 33: 78 (key); West-Eberhard et al., 1995. Hym. Costa Rica: 574. Misidentification]. I use this name in order to commemorate Bertoni ’s memory, who labeled with it a female specimen that perfectly fits with the key and description given by Bohart and Stange (1965) for their concept of Zethus costarricensis. Holotype female (Figs. 1, 2): Length from antennal insertions to apex of tergum 1997 DISCOELIUS STRIGOSUS COSTARRICENSIS 217 1 Fig. 1. Zethus smidtianus n. sp., female habitus. 218 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) II about 13 mm. Black with yellow as follows: subapical and lateral marks on clyp- eus, scrobal spots, humeral spots, anterior and inner posterior comers of the tegulae, parategulae, a pair of oval transversal spots on metanotum, subapical bands on terga I-II and sternum II, longitudinal anterior line on fore and mesotibiae. Wings hyaline yellowish with light brownish infuscation along the costal region of the forewing. Venation dark brown. Clypeus striatopunctate, frons logitudinaly, rather irregularly striate an with strong punctures into the striae, ocular sinus striatopunctate, vertex and gena with strong punctuation, pronotum strongly punctate, some longitudinal wrinkles present on the subhumeral area, mesoscutum longitudinaly, rather irregularly striate and with punc- tuation more casual than on frons, disk of scutellum flat, polished and with large, sparse, rather shallow punctures and very rare almost absent micropunctation on the intervals, upper face of metanotum sculptured as scutellum, posterior face of meta- notum without macropunctures, upper and posterior face of propodeum with coarse reticulation that obscures the transversal striae that otherwise are well developed towards the middle, lateral face of the propodeum with large, very sparse punctures that progressively concentrate towards the anterior comer, mesopleura strongly punc- tate, tergum I polished and with strong punctuation except for the subapical band, that lacks punctures, tergum II moderately punctate and with macropunctures con- centrated on a polished subapical band, this last having a very narrow impunctate apical region followed by a narrow crenulated furrow preceding the apical lamella, which is slightly raised, tergum III subapically with macropunctures finer than those on tergum II, almost immediately followed by a very narrow crenulated furrow that precedes the apical lamella, this one not raised at apex. Head, mesoscutum and tergum I with pale, sub-erect, sparse pilosity, specially sparse on the latter, remaining terga with fulvous, decumbent, relatively dense pi- losity, denser on tergite II. Metanotum and mainly propodeum with pale, decumbent, relatively dense pilosity as well as suberect pilosity. Clypeus truncated apically, interantennal carina well developed and entering into the clypeus, OOL: POL = 3 : 2, 2, notauli almost complete, propodeum with a distinct medial longitudinal carina, tergum I in dorsal view suddenly expanding after the stem and more or less parallel sided, subapical polished band longer submedially, here it is longer than the apical band, this last being longer than a midocellar di- ameter, midtibia with two spurs, the hind one being a bit longer. Male (Figs. 3, 4, 5, 6, 7, 8 and 9): Similar to the female. Length from antennal insertion to the apex of tergum II about 8 mm. Yellow color on clypeus forming a wide U-shaped mark, also yellow are a mandibular spot and the inner apex of fla- gellomere VIII, as well as almost the whole inner face of flagellomeres IX and X (in the UCD male there is not yellow on flagellomere VIII and flagellomere IX is yellow only at apex). The punctuation on frons and mesoscutum is more evident than in the female, being the sculpture almost striatopunctate. Clypeus with an apical notch. Genitalia similar to those figured by Bohart and Stange (1965: 201, figs. 304 y 305) for Z. evansi, but the mast (“asta” following Giordani Soika, 1978: 7, fig. 1) of the cuspis lacks pilosity on the lower margin and the apex of the basivolsella is more expanded. Holotype: Costa Rica: San Jose, 3168, female, Bertoni Collection, IBNP (I labeled it with the reference number “A: 41” as part of the recovery work). 1997 DISCOELIUS STRIGOSUS COSTARRICENSIS 219 Figs. 2-9. Zethus smidtianus n. sp. 2. female. 3-9. male. 2 and 4. Head in frontal view. 3. Last antennal segments. 5. Paramere and volsella in inner profile view. 6. Detail of the volsella. 7. Ventral view of paramere and volsella. 8. Ventral view of the aedeagus. 9. Profile view of the aedeagus. 220 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Paratypes: Guatemala: Sacatepequez, Capetillo, 5000 ft. 20. viii. 1974, 1 female (C. y P. Vaurie cols., F. Johnson leg.), UCD. Costa Rica: San Jose, 1913, 1 male, UCD; San Jose, San Antonio de Escazu, 1300 msnm, 1 male, AMNH. There are additional specimens known (Bohart and Stange, 1965) but I was unable to examine: Colombia: Cauca, Villa Elvira, 1 female, Giordani Soika Collection; Costa Rica: San Jose, 1 female, UCD; San Jose, 1 female (M. Valerio col.), USNM. According to Menke (in litt.), who sent to me the complete data, this later specimen seems to be conspecific with photographs of the holotype I sent to him. ACKNOWLEDGMENTS Eor the loan of specimens I thank Lynn S. Kimsey and S. L. Heydon of the University of California, Davis, USA (UCD) and James M. Carpenter of the American Museum of Natural History, New York, USA (AMNH). I thank this last as well as Arnold S. Menke of the United States National Museum of Natural History, Washington, USA (USNM), Abraham Willink of the Institute Miguel Lillo, Tucuman, Argentina (IML), Carlo Dlouhy of the Universidad Na- cional de Asuncion (UNA) and John A. Kochalka of the Museo Nacional de Historia Natural del Paraguay, San Lorenzo, Paraguay (IBNP) for the comments and continuous encourage regarding the recovery of Bertoni collection. LITERATURE CITED Bertoni, A. W. 1925. Himenopteros nuevos o poco conocidos. Rev. Soc. Cient. Paraguay. (2) 1:74-79. Bohart, R. M. and L. A. Stange. 1965. A revision of the genus Zethus Eabricius in the Western Hemisphere (Hymenoptera: Eumenidae). Univ. Calif. Pub. Entom. 40:1-208. Cameron, P. 1907. On some new Central American Vespidae. Entomologist 40:62-64, 79-83. Garcete Barrett, B. R. 1996. The collection of Arnaldo de Winkelried Bertoni. Sphecos 30:25- 26. Giordani Soika, A. 1978. Revisione degli eumenidi neotropical! appartenenti ai generi Eumenes Latr., Omicron (Sauss.), Pararaphidoglossa Schulth. ed affini. Boll. Mus. Civ. Venezia, XXXIX:7-420. International Commission on Zoological Nomenclature. 1985. International Code of Zoological Nomenclature, 3rd ed. International Trust for Zoological Nomenclature, London, xx + 338 pp. Saussure, H. de. 1975. Synopsis of American Wasps. Smithsonian Misc. Coll. 254, 393 pp. Stange, L. A. 1978. Los Zethus del desierto costal de Peru. Acta Zool. Lilloana 33:71-78. West-Eberhard, M. J., J. M. Carpenter and P. E. Hanson. 1995. The Vespid Wasps (Vespidae). Ch. 15, pp. 561-587 in P. E. Hanson and I. D. Gauld (eds.). The Hymenoptera of Costa Rica. Oxford University Press, Oxford, xx + 893 pp. Willink, A. 1982. Himenopteros neotropicales. Su origen, ecologia, comportamiento y distri- bucion. Zoologia Neotropical. Actas del VIII Congreso Latinoamericano de Zoologia. Ed. Pedro Salinas. 1:71-90. Zavattari, E. 1912. Materialen fiir eine Monographie der Neotropischen Eumeniden. Archiv fiir Naturgeschichte 78 (Abt. A), Heft 1:87-118. Received and accepted 28 October 1997. J. New York Entomol. Soc. 105(3-4):22 1-229, 1997 OBSERVATIONS ON PHENOLOGY, DEVELOPMENT, AND MORTALITY OF LARVAE OF THE HAZELNUT WEEVIL {CURCULIO OBTUSUS (BLANCHARD): CURCULIONIDAE) IN NUTS OF BEAKED HAZELNUT (CORYLUS CORNUTA MARSHALL: BETULACEAE) IN THICKETS IN MAINE' L. W. Treadwell^ and R. H. Storch Department of Biological Sciences, University of Maine, Orono, Maine 04469 Abstract. — Little is known about the biology of the hazelnut weevil Curculio obtusus (Blan- chard), though closely-related species are widespread and well studied. The objective of this study was to document details of development of larvae in nuts, including mortality factors, assessment of the infrequent phenomenon of multiple infestation of nuts, and confirmation of 4 larval instars. Samples of beaked hazelnuts taken every 4 days over the course of the summer, 1995, revealed the development of larvae through 4 instars, beginning with eggs in early June and progressing through 4th-stage grubs exiting nuts in late August. At the peak of infestation, in early August, 71% of the sampled nuts were infested with weevil larvae. Decay and non- formation of nut kernels accounted for an estimated 10% mortality among the larvae. Multiple infestation accounted for an additional 8% mortality due to interference competition. Nut pre- dation also caused mortality of an estimated 10% of larvae. The genus Curculio (Curculionidae; Curculioninae) is comprised of 27 species of nut- and acom-infesting weevils in North America (Gibson, 1969). Most (23 species) inhabit oak trees (Quercus spp.); only C. obtusus, the hazelnut weevil, is known to specialize in hazelnuts (Corylus spp.) in the eastern half of the United States (Ham- ilton, 1890; Brooks, 1910; Gibson, 1969, 1985a, 1985b). This weevil exploits the nuts of wild {Corylus americana Walter) and beaked (C. cornuta Marshall) hazelnut not only for food at all stages but also for oviposition sites (Gibson, 1969). Adult Curculio species emerge from the soil in early summer and fly to the nearest host tree or shrub where they feed on immature nuts or acorns and mate (Gibson, 1969; Raney and Eikenbary, 1968). The species are dimorphic; the females’ rostra are somewhat to much longer than those of males, depending on species, and are used to excavate egg chambers in the nuts as well as to feed (Brooks, 1910; Gibson, 1969). The vermiform Curculio grubs are reported to pass through 4 instars feeding inside their host nuts, destroying the nut kernel in the process. At maturity in the fall, they chew an exit hole in the nutshell and drop to the ground where they immediately burrow almost straight down, to depths from 9 to 28 cm among pecan weevils (C. ' Based on data submitted by L. W. Treadwell in partial fulfillment of the requirements for the MS degree in Entomology, University of Maine, Orono. 2 Present address: Department of Entomology and Nematology, University of Florida, Gaines- ville, Florida 32611. 222 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) caryae [Horn]) (Gibson, 1969; Harris, 1975; Harp and Van Cleave, 1976; Harrison et al., 1993), and enter a state of diapause. Curculio species pass the majority of their lives as subterranean diapausing larvae. The length of diapause varies within species; Menu and Debouzie (1993) found, for example, that up to 4 years could be spent underground by a small proportion of chestnut weevil (C. elephas Gyllenhal) grubs in France; 32-56% emerged after 2 to 3 years. The few references to the hazelnut weevil C. obtusus consist mainly of early naturalist descriptions (e.g., Hamilton, 1890; Brooks, 1910; Hutchings, 1927). Few details have been reported about its biology, though its host species range from Quebec to Georgia and westward into the midwestem U.S. (Femald, 1950; Gibson, 1969). Furthermore, its host species have at times been considered to have com- mercial potential, particularly in Canada (e.g., Hutchings, 1927; St. Pierre, 1992). Curculio obtusus larvae develop singly in a nut, unlike many of their congeners. Vrabl et al. (1979, p. 363) assert, of the European hazelnut weevil C. nucum L., that <4 eggs can be found in the shell of cultivated hazelnuts {Corylus avellana L.) in Slovenia, “but into the fruit itself there creeps only 1 larva.” Multiple infestation, which occurs at a low level among C. obtusus (Gibson, 1969), is maladaptive; as with C. nucum, only 1 larva will succeed in nuts containing >1 larva (pers. obs.). Interior galls and other deformities in the nutshell are caused by this species in wild and beaked hazelnuts and also by the European hazelnut weevil in cultivated hazelnuts (Rabaud, 1913; Gibson, 1969; Meyer, 1987). Curculio obtusus, unlike the other Curculio species, has been reported to pass 2 instars feeding in the soft shell and shell lining before passing into the kernel (Gibson, 1969). The objective of the current study was to investigate and document details of larval development of the hazelnut weevil, including an assessment of the phenom- enon of multiple infestation, determination of mortality factors, and confirmation of the number of larval stages. These observations were part of a larger study that included documentation of adult behavior and intershrub movements detailed by Treadwell (1996). MATERIALS AND METHODS Study site and host plant descriptions: The study was conducted in a secondary succession of woodland in T32 MD, Hancock County, Maine (44°58'N, 68°27'W), at an elevation of ca. 82 m, on land owned by Champion International Corporation and minimally maintained for primitive camping. The plant community consisted of a grey birch (Betula populifolia Marshall)-beaked hazelnut-blueberry (Vaccinium angustifolium Alton) sere on sandy soil. The site was chosen because of a profusion of “brush-stage” (Hsiung, 1951) hazelnut thickets and observations of a high degree of hazelnut weevil activity in the area the previous summer. Beaked hazelnut, so-called for the projecting involucre of united bracts which enclose its nut, is a monoecious shrub distributed throughout woodlands and forests of central North America. It is one of only 3 species of Corylus native to North America and is grouped by some in the hazel family, Corylaceae, along with Ostrya Scopoli and Carpinus L. (ironwood), Betula L. (birch), 2iwd Alnus B. Ehrhart (alder) (Femald, 1950). Others place Corylus in the family Betulaceae (Hsiung, 1951). It sprouts vigorously from underground modified stems and, in New England and the 1997 HAZELNUT WEEVIL 223 Adirondacks, is one of the 6 most common shrub-stratum species in successional progressions from overgrown pastures to climax forests of hemlock (Tsuga cana- densis [L.] Carriere.), beech (Fagus grandifolia F. Ehrhart), and sugar maple (Acer saccharum Marshall) (Hsiung, 1951). In situations of high light intensity, found by Hsiung (1951) to be the critical factor in density of Corylus cornuta, the shrub attains maximum vegetative reproduction and can appear as dense thickets. Larval development of the hazelnut weevil: Samples of 75->100 nuts (x = 103 ±18 nuts, N = 22 samples) were collected haphazardly from thickets every 4 days from 12 June to 4 September 1995 for dissection, to monitor larval development as well as progression of nut damage. Nuts were transported on ice and immediately stored in a freezer until dissection under a microscope within 3 days. Dissected nuts were categorized as (a) intact (no damage to nut meat), (b) no kernel (shell fully formed but containing only a cottony lining), (c) decayed, (d) weevil-infested (con- taining eggs, larvae, or exit hole), and (e) damaged but not infested (i.e., outwardly punctured and/or deformed). Weevil eggs and larvae at each of 4 stages found in dissected nuts were counted and preserved in 70% ethanol. Twenty-four individuals of each of the 4 assumed larval stages were selected randomly and the width of their head capsules measured with an ocular micrometer in a dissecting microscope. RESULTS A total 2,269 nuts were collected and dissected between 12 June and 4 September 1995 at 4-day intervals. Of these, 873, or 38.5%, were infested with weevil eggs or larvae, with a peak 70.6% infestation occurring on 7 August (Fig. la). Decayed nuts peaked at 30.4% on 18 July (Fig. lb). Eggs were first found in the layers of nutshells on 20 June (Fig. 2), 17 days after the first adults were observed and 2 days after adults were first observed mating (Treadwell, 1996). First instars were found feeding in nut shell layers or interior galls beginning on 6 July, about 2 weeks after the onset of oviposition. Fourth instars were found beginning on 7 August, when percentage of total nuts infested with this stage suddenly increased from 0 to >17%. Exit holes were first observed in the 11 August sample. Complete larval development, from egg to exit, thus takes about 36 days. Weevils had emerged from 71.4% of the infested nuts in the 4 September sample. Sampling was discontinued at this date because predation on nuts, presumably by the many red squirrels (Tamiasciarus hudsonicus gymnicus [Bangs]) and chipmunks (Tamias striatus L.) in the area, had become noticeably heavy on 27 August. Empty shells were more and more frequently found strewn under the shrubs and thickets, and natural nut drop had also begun by this date; thus nuts for sampling were difficult to find. Additionally, the proportion of nuts with no kernel increased from ca. 20% on 27 August to almost 40% on 4 September (Fig. lb). Multiple infestation occurred in 70 nuts, or 8% of the total 873 infested, and peaked on 30 July, when 22.6% of infested nuts contained >1 larva (Fig. la). The majority of multiple inhabited nuts contained 2 larvae, though 10.4% contained 3 larvae and a single nut contained 4 larvae (Fig. 3). In only 2 cases was a 4th-stage larva found in association with another larva. In one of these, surprisingly, the 4th instar was found partially consumed by a much smaller 2nd instar. 224 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Fig. E Hazelnut weevil larval infestation, beaked hazel nut conditions, and climatic con- ditions at T32 MD, Hancock County, Maine, 28 May-4 Sept. 1995. (a) Percent of nuts infested, and percent of infested nuts containing >1 larva, in samples collected at 4-day intervals; (b) percent punctured/damaged, decayed, and no-kernel nuts in samples collected at 4-day intervals; (c) daily rainfall, solar radiation, and average air temperatures. precipitation (mm) 1997 HAZELNUT WEEVIL 225 < Eig. 2. Percent of larval weevil instars among sequential samples of beaked hazelnuts, 20 June to 4 Sept. 1995, T32 MD, Hancock County, Maine. Eggs appeared most often in the top Vs or Va of the shell. First instars were observed to feed first within the shell layers, sometimes within the exaggerated thickness of a gall, until breaking through to the shell lining. They then fed in the lining in a fairly straight line down to the bottom of the nut. Entry into the kernel was made at a distinctive round hole chewed through the thin skin, and ecdysis to the 2nd stage occurred soon after this. There was no evidence for feeding in the shell lining by 2nd instars. In the early stages, multiple larvae could easily be found by looking for multiple entry holes into the kernel. Sometimes, however, one larva could be tracked between holes connected by longitudinal ruts in the lining; some would apparently sample the nut kernel before settling down to feeding. Mean head capsule widths among the 4 stages appeared to vary only slightly from the predictions of Dyar’s Law (Dyar, 1890); i.e., a constant ratio (in this case ca. 0.60) between the head capsule widths of each successive stage (Fig. 4). C3w/4 larvae □ w/3 larvae ■ w/2 larvae ir> ro -t- T- CM CO Fig. 3. Proportions of infested nuts containing 2, 3, or 4 larvae. 226 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-1) Eig. 4. Mean width of larval head capsules at 4 stages (N = 24/stage). DISCUSSION Anomalous trends in the data: A peak and then decline in proportions of infested nuts seems counter-intuitive: the proportion of nuts infested by weevils should level off at some point, since the nuts do not renew themselves and infestation is a per- manent state. Pucci (1992: 9) reported a similar trend in data for European hazelnut weevils C. nucum infesting commercial hazelnuts in Italy, and attributed it to “the fruit drop effect”: hazelnuts with larval exit holes tended to drop earlier than un- infested nuts. A similar phenomenon among beaked hazelnuts infested by C. obtusus would mean that the chances of picking an uninfested nut improve after a peak of infestation in early August, though the chances of its being decayed or unfilled are also greater (Fig. la). Probably these latter are lighter, lack abscissive mechanisms (why waste resources on an unfruitful shell?), and are instinctively passed over by nut predators; thus they will appear in greater proportions at the end of the season. Two reasons might be given for the large proportion of 1st instars found in the sequential samples of nuts (Fig. 2). First, because eggs were deposited in shell layers it is likely that an unknown number were overlooked in dissection. An effort was made to examine all punctures and wounds in the shell, but eggs were observed to rupture and spill extremely easily, so some “disappeared” in the dissection process; numbers of eggs are most likely under-reported. For these reasons, occurrence of >1 egg is no doubt also under-reported. Secondly, some first instars lingered into the final sample, a full 8 weeks after this stage had first appeared. An extended first larval stage would cause an exaggeration of their abundance proportionate to the other stages; members of the same cohort would be counted >1 time. Lowered quality or availability of nutrients has been shown to inhibit ecdysis in many insect species. Sehnal (1985) found, for example, that caterpillars of the greater wax moth Galleria mellonella (L.) molted after 15 hours on a normal diet but required 40 hours when the diet was mixed equally with sawdust. Release of prothoracicotropic neurohormone, which stimulates release of molt-inducing ecdysteroid, has been shown to be dependent on attainment of a cer- tain “critical” body size increment (Sehnal, 1985). 1997 HAZELNUT WEEVIL 227 First instars in the last few samples were all found in decaying nuts or nuts with no kernel. Apparently the cottony parenchyma of no-kemel nuts can sustain the life of a weevil larva but does not provide sufficient nutrients or appropriate cues for ecdysis. Besides obviously decaying and unfilled nuts, many “normal-appearing” kernels might have been affected by some factor, such as the long period of no rain as nuts were forming (Fig. Ic), which lowered their quality enough to inhibit ecdysis. Factors in mortality of larvae: Conditions of unfilled nuts and decay appear to be major mortality factors among young weevil larvae. Females unwittingly deposit eggs into shells in which kernels do not form and/or into which fungi and myceto- philous flies enter, in many cases, ironically, through the weevil oviposition puncture (Winston, 1956), and initiate a decay process. Larvae hatch into an environment unsuitable for growth and development but many subsist for weeks before dying of desiccation or starvation. Ten percent of the nuts in sequential samples over the season showed evidence of rot or decay, with or without ubiquitous, unidentified microscopic maggots — from 1 to 15 in any one nut. Winston (1956) reported Fusarium sp. and Penicillium sp. responsible for initiating decay in acorns of red oak (Quercus rubra L.) in Illinois. He also found maggots, identified as Mycodiplosis sp. and Rubsoamenia sp. (Ceci- domyiidae), to be spreading spores of the fungi. As in the case of maggots observed in hazelnuts, they created areas of decay around themselves; weevil larvae in mag- got-infested nuts are certain not to survive. An unconscious sampling bias might have been introduced against decayed nuts in the field because of their often obvious appearance. A more accurate estimate of their effect on larval populations can be gained by analyzing an absolute sample of nuts (N = 1,246) collected from 33 discrete beaked hazelnut shrubs on 3 August 1995 (Treadwell, 1996). Here 15.3% of the total were in a state of decay that ren- dered the nut kernel unfit for weevil sustenance. An accurate count of larvae in decaying nuts could not be kept because the cadavers were also broken down by invading hyphae and maggots. Decay and no-kemel conditions together claim ca. 25% of a hazelnut crop. If 40% of those nuts are weevil-infested (Treadwell, 1996), an estimated 10% of total larvae do not survive because of pathological nut condi- tions. Since it appears that ca. 30 days are required to complete larval development after ecdysis into the 2nd instar, gmbs that were still at 1st instar on 15 August, 9-10% of the larval population, most likely did not survive. Quality of mature nuts can be assumed to be quite different from that of young developing kernels with which early instars normally coincide. Additionally, the fate of the hazelnut itself became more and more tenuous as the summer came to an end. As mentioned, red squirrels and chipmunks were observed at this site and were noted by Hsiung (1951) to consume or bury “the great proportion of the [beaked] hazelnut crop” at a Minnesota study site. It is not known whether these two rodents have any preference for uninfested nuts. Grey squirrels {Sciurus carolinensis Gme- lin), however, have been observed to detach and sniff or taste European hazelnuts infested with C. nucum before discarding them (Lloyd, 1968). White-footed mice {Peromyscus leucopus Thomas) prey on detached acorns of white oak {Q. alba L.) with no preference for those infested or uninfested by larvae of the weevils Curculio pardalis (Chittenden) and Conotrachelus naso LeConte (Semel and Andersen, 1988). 228 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) Additionally, detached nuts were observed in the laboratory either to desiccate or to start decaying within a few days. Multiply-infested nuts account for another 8% of larval mortality. These represent larvae that were selected out by a critical interference competition between nut- mates. Since multiple larvae reached levels of >20% of infested nuts in the sequen- tial random samples, a competitive “edge” inside the nutshell might seem a measure of fitness. In the similar case of the cowpea weevil Callosobruchus maculatus (Fabricius), small beans can sustain only 1 larva to maturity. Among the larva of this species, in any bean regardless of size one larva will out-compete any others in survival. Mitchell (1983) speculates that interference competition, “activity which directly or indirectly limits a competitor’s access to a resource” (Prokopy et al., 1984, p. 307), might have evolved to ensure the success of at least one larva, rather than a situation of exploitation competition in which both would die if consuming a small bean equally until it was depleted. For the hazelnut weevil there can be no doubt as to the optimum competition strategy; 1 larva can and usually does consume an entire kernel. In summary, then, these observations suggest almost 30% mortality among nut- infesting hazelnut weevil larvae due to decay, interference competition, and nut predation. Conclusions: The inside of a hazelnut seems a relatively secure, insulated habitat for growth and development. At the scale of a weevil, however, this environment presents a variety of hazards to which the insect has adapted and evolved. Evidence of parasitization of nut-inhabiting larvae was not detected, but rearing of larvae extracted from nuts, in a controlled environment, would reveal whether parasitization has taken place. Additionally, a longer-term study would shed some light on ques- tions regarding synchronicity of life cycle with host plant phenology. ACKNOWLEDGMENTS The field assistance of Bancroft Whitely is gratefully acknowledged. This study was sup- ported, in part, by the Maine Agricultural and Forestry Experiment Station and The Graduate School of the University of Maine. This article is published as Maine Agricultural and Forestry Experiment Station Publication MAFES No. 2114. LITERATURE CITED Brooks, F. C. 1910. Snout beetles that injure nuts. W. Va. Agric. Expt. Sta. 128:145-185. Dyar, H. G. 1890. The number of molts of lepidopterous larvae. Psyche 5:420-422. Fernald, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Co., New York. Gibson, L. P. 1969. Monograph of the genus Curculio in the New World (Coleoptera: Curcu- lionidae). Part I: United States and Canada. Misc. Publ. Entomol. Soc. Am. 6(5):239- 285. Gibson, L. P. 1985a. Catalog of the Coleoptera of America north of Mexico. Family: Curcu- lionidae. Subfamily: Curculioninae. USDA-ARS Agriculture Handbook no. 529- 143b, 14 pp. Gibson, L. P. 1985b. Description and key to larvae of Curculio spp. of eastern United States and Canada (Coleoptera: Curculionidae). Proc. Entomol. Soc. Wash. 87(3):554-563. Hamilton, J. 1890. Balaninus — its food habits. Can. Entomol. 22(1): 1-8. 1997 HAZELNUT WEEVIL 229 Harp, S. J. and H. W. Van Cleave. 1976. Biology of the subterranean life stages of the pecan weevil in two soil types. Southwest. Entomol. l(l):31-34. Harris, M. K. 1975. Pecan weevil distribution in some Texas soils. Environ. Entomol. 4(5); 849-853. Harrison, R. D., W. A. Gardner, W. E. Tollner and D. J. Kinard. 1993. X-ray computed tomog- raphy studies of the burrowing behavior of fourth-instar pecan weevil (Coleoptera: Cur- culionidae). J. Econ. Entomol. 86(6): 17 14-17 19. Hsiung, W.-Y. 1951. An ecological study of beaked hazel (Corylus cornuta Marsh.) in the Cloquet Experimental Forest, Minnesota. Ph.D. thesis. University of Minnesota. 117 pp. Hutchings, C. B. 1927. A study of Balaninus obtusus Blanchard; or, a life history in a hazel nutshell. Entomol. Soc. Ontario Annu. Rep. 57:9-12. Lloyd, H. G. 1968. Observations on nut selection by a hand-reared grey squirrel (Sciums carolinensis). J. Zool. Proc. Zool. Soc. London 155:240-244. Menu, E and D. Debouzie. 1993. Coin-flipping plasticity and prolonged diapause in insects: example of the chestnut weevil Curculio elephas (Coleoptera: Curculionidae). Oecologia 93:367-373. Meyer, J. 1987. Plant Galls and Gall Inducers. Gebriider Borntraeger, Berlin, viii + 291 pp. Mitchell, R. 1983. Effects of host-plant variability on the fitness of sedentary insects. In R. E Denno and M. S. McClure (eds.). Variable Plants and Herbivores in Natural and Managed Systems, 343-370. Academic Press, New York. Prokopy, R. J., B. D. Roitberg and A. L. Averill. 1984. Resource partitioning. In W. J. Bell and R. T. Carde (eds.). Chemical Ecology of Insects. Sinauer, Sunderland, MA. Pucci, C. Studies on population dynamics of Balaninus nucum L. (Col., Curculionidae) noxious to the hazel {Corylus avellana L.) in Northern Latium (Central Italy). Zeit. Ange. En- tomol. 114:5-16. Rabaud, E. 1913. La cryptocedicie du ver des noisettes {Balaninus nucum L.) et la signification biologique des galles. C. R. Acad. Sci. Paris 156:253-255. Raney, H. G. and R. D. Eikenbary. 1968. Investigations on flight habits of the pecan weevil, Curculio caryae (Coleoptera: Curculionidae). Can. Entomol. 100:1091-1095. Sehnal, E 1985. Growth and life cycles. In G. A. Kerkut and L. I. Gilbert (eds.). Comprehensive Insect Physiology, Biochemistry and Pharmacology, Vol. 2, 1-86. Pergamon, Oxford. Semel, B. and D. C. Andersen. 1988. Vulnerability of acorn weevils (Coleoptera: Curculionidae) and attractiveness of weevils and infested Quercus alba acorns to Perornyscus leucopus and Blarina brevicauda. Amer. Midi. Nat. 1 19(2):385-393. St. Pierre, R. G. 1992. The development of native fruit species as horticultural crops in Sas- katchewan. HortScience 27(8):866, 947. Treadwell, L. W. 1996. Aspects of larval development, intershrub movement, and nut infestation by the hazelnut weevil {Curculio obtusus) utilizing discrete shrubs of beaked hazelnut {Corylus cornuta). MS thesis. University of Maine. 58 pp. Vrabl, S., K. Beber and G. Matis. 1979. A contribution to the biology and noxiousness of nut weevil {Curculio nucum L.). Plant Protection (Zastita Bilja) 30(4):357-364. Winston, P. W. 1956. The acorn microsere, with special reference to arthropods. Ecology 37(1): 120-132. Received 15 October 1996; accepted 3 September 1997. J. New York Entomol. Soc. 105(3-4);230-235, 1997 SENSORY STRUCTURES OF THE ANTENNAE OF NANNOTRIGONA TESTACEICORNIS (APIDAE: MELIPONINAE)’ Antonio Carlos Stort and Monica M. B. Morals- Alves Department of Biology, Institute of Biosciences, UNESP, 13.506-900 Rio Claro, SP, Brazil Department of Exact and Biological Sciences, University Center of Dourados, UFMS, 79.825-070 Dourados, MS, Brazil. Abstract. — The total number and distribution per antennal flagellomere of sensilla placodea (olfactory disks), sensilla coeloconica, sensilla ampullacea and sensilla campaniformia were determined in workers of Nannotrigona testaceicornis Lepeletier a stingless bee species quite common in Brazil. The distribution of the sensilla was uniform, with the largest number oc- curring in flagellomere 10 and gradually decreasing in the direction of the basal flagellomeres in a way similar to that observed in Scaptotrigona postica Latreille. Nannotrigona testaceicornis had a larger number of sensilla ampullacea and a smaller number of sensilla coeloconica and sensilla campaniformia than Scaptotrigona postica. Although Nan- notrigona testaceicornis does not communicate through the formation of pheromone trails, this species presents a larger quantity of sensilla placodea (relative to the length of the flagellum) than Scaptotrigona postica. Nannotrigona testaceicornis, a stingless bee species quite common in Brazil, is found from the north of Parana up to Mexico (Nogueira Neto, 1970). This species presents on average populous colonies, generally consisting of 2,000 to 3,000 indi- viduals (Lindauer and Kerr, 1960). Nannotrigona testaceicornis has been studied from different aspects such as ovi- position behavior (Sakagami and Zucchi, 1966), communication (Kerr and Esch, 1965), glandular system (Cruz-Landim, 1967), taxonomy (Moure, 1951), nest struc- ture (Nogueira Neto, 1970) and morphometry (Cunha, 1973). On the other hand, only little information exists with respect to the antennal sensilla placodea of the workers of this species (Johnson and Howard, 1987). In order to expand the knowledge about the outer morphology of Brazilian sting- less bees, we determined the number and distribution of sensilla placodea, sensilla coeloconica, sensilla ampullacea and sensilla campaniformia of the antennae of Nan- notrigona testaceicornis workers. MATERIAL AND METHODS Samples of Nannotrigona testaceicornis Lepeletier workers were collected from colonies maintained at the Animal House of the Biosciences Institute of Rio Claro. The bees were anesthetized and killed in an ether chamber, fixed in modified Kar- novsky (2% glutaraldehyde and 2% paraformaldehyde in 0.1 M phosphate buffer, pH 7.2) for 24 hours and stored in 70% alcohol. The antennae were separated from Research supported by CNPq and FAPESP. 1997 ANTENNAE OE NANNOTRIGONA TESTACEICORNIS 231 the head, treated with ultrasound for 1 minute to remove dirt particles and glued to a metal support. The antennae were then sputtered with a thin gold layer using an Edwards Sputter apparatus, model S150B, and observed with a model T330A Jeol scanning electron mi- croscope of the Chemistry Institute, UNESP, Araraquara. For sensilla placodea (olfactory disks) counts, the antennal flagellomeres were separated and opened with the aid of two entomology pins and mounted on balsam between a slide and a coverslip. Each flagellomere was photographed with a Zeiss photomicroscope II, and the film was projected onto a paper screen where the sen- sory structures were counted. The ratio number of sensilla placodea to length of the flagellum was also studied. The length of the flagellum was measured under a dissecting microscope equipped with an optical micrometer. Ten worker bees were used for the observations and counts. RESULTS In flagellomere 10 the set of sensilla campaniformia was located above the sets of sensilla coeloconica and sensilla ampullacea (Fig. lA), with these two types of sensilla (coeloconica and ampullacea) being clearly distinguishable. The total number of sensilla ampullacea in Nannotrigona testaceicornis workers was larger than that of sensilla coeloconica, with the largest number of these sensilla and of sensilla placodea occurring in flagellomere 10 and decreasing in the direction of the most basal flagellomeres (Table 1). The set of sensilla campaniformia was only observed in flagellomere 10, having on average 5.800 ± 0.421 units (Table 1). No set of these sensilla was observed in the other flagellomeres below flagellomere 10 of the antennae of Nannotrigona tes- taceicornis workers (Fig. IB), and an isolated sensilla campaniformia rarely oc- curred. DISCUSSION The location of the set of sensilla campaniformia above the sets of sensilla coe- loconica and sensilla ampullacea in flagellomere 10 of the antennae of Nannotrigona testaceicornis workers is similar to that observed in Scaptotrigona postica (Stort and Barelli, 1981; Stort and Moraes- Alves, 1997) and differs from what occurs in Apis mellifera where the set of sensilla campaniformia is located laterally to the set of sensilla coeloconica + ampullacea (Dietz and Humphreys, 1971; Stort and Moraes- Alves, 1997). The morphological differences between sensilla coeloconica and sensilla ampul- lacea observed in Nannotrigona testaceicornis are similar to those observed in Scap- totrigona postica (Stort and Barelli, 1981). Nannotrigona testaceicornis workers possess a smaller total number of sensilla coeloconica and a larger total number of antennal sensilla ampullacea (Table 1) than Scaptotrigona postica (Stort and Moraes- Alves, 1997). Regarding the total sum of these two types of sensilla (sensilla coeloconica + sensilla ampullacea), Nannotrigona testaceicornis presents a smaller quantity (mean: 48.100 ± 5.606) than Scaptotrigona postica (mean: 52.800 ± 5.266). According to 232 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) Fig. 1. Scanning electron microscopy micrographies of the antennal flagellomeres of Nan- notrigona testaceicornis worker. A — flagellomere 10 showing sensilla campaniformia (long arrow), sensilla ampullacea (short arrow), sensilla coeloconica (median arrow) and sensilla placodea (S P). B — flagellomere 9 showing sensilla ampullacea (short arrow), sensilla coelo- conica (long arrow) and sensilla placodea (S P). 1997 ANTENNAE OE NANNOTRIGONA TESTACEICORNIS 233 Table 1. Means (± standard deviations) of the number of sensilla placodea, sensilla coelo- conica, sensilla ampullacea and sensilla campaniformia of the antennal flagellomeres (E) of Nannotrigona testaceicornis workers. Sensilla placodea Sensilla coeloconica Sensilla ampullacea Sensilla campaniformia F,o 145.600 + 5.274 4.300 H- 0.823 6.800 -1- 0.918 5.800 ± 0.421 F9 104.800 ± 4.638 3.900 -h 0.994 5.400 H- 1.577 — Fg 103.600 -H 4.247 3.500 -h 0.707 4.300 H- 2.057 — F7 87.300 4.762 3.400 -h 0.843 3.100 -1- 0.994 — F6 87.500 + 3.719 2.300 -h 1.159 3.200 -H 1.988 — F5 76.800 7.656 1.800 -h 0.788 1.900 H- 0.737 — F4 76.700 + 4.990 1.700 ~h 0.483 1.800 -H 1.135 — F3 64.300 -H 4.243 0.400 -h 0.699 0.300 -h 0.483 — F2 48.200 -H 4.825 — — — Total 794.800 22.592 21.100 -f- 2.960 27.000 -h 4.876 5.800 ± 0.421 Kuwabara and Takeda (1956), these structures are hygroreceptor organs, a fact that was confirmed by Lacher (1964). Nannotrigona testaceicornis would therefore be less sensitive in detecting the degree of environmental humidity than Scaptotrigona postica. In contrast. Africanized Apis mellifera, which present on average 71.900 ± 7.311 sensilla coeloconica + ampullacea (Stort and Rebustini, 1997), appear to be more sensitive in detecting humidity than these two stingless bee species. Both Nannotrigona testaceicornis and Scaptotrigona postica have one set of sen- silla campaniformia in flagellomere 10 and rarely present other sensilla of this type, even isolated, together with sensilla coeloconica and sensilla ampullacea in the fla- gellomeres below number 10. Nannotrigona presents a smaller quantity of sensilla campaniformia (Table 1) than Scaptotrigona (Stort and Moraes- Alves, 1997). Therefore, this species should be less sensitive to temperature and CO2 since these sensilla are related to the perception of these environmental parameters (Dietz and Humphreys, 1971). According to Stort and Rebustini (1997), Africanized Apis mellifera workers have a larger quantity of sensilla campaniformia (mean: 18.700 ± 2.710) since these sensilla are present in all antennal flagellomeres, and this species may therefore be more sensitive to these environmental factors than the two stingless bee species mentioned here. The total number of sensilla placodea (olfactory disks) per antenna in Nannotri- gona testaceicornis was on average 794.800 ± 22.592 (Table 1), a number 30.397% smaller than that observed in the antenna of Scaptotrigona postica (Stort and Barelli, 1981) and 327.126% smaller than that observed in Africanized Apis mellifera (Stort, 1979). The total number of sensilla placodea of Nannotrigona testaceicornis obtained in the present study disagrees completely with the number (1,858 ± 338) found by Johnson and Howard (1987). The total number of sensilla placodea, however, when considered in relation to the length of the antenna which was determined by calculating the number of sensilla placodea/flagellum length ratio, showed a difference. The ratios obtained were 24.380 for Nannotrigona, 24.102 for Scaptotrigona and 39.877 for Africanized Apw. This means that Nannotrigona presents, proportionally to the size of the flagellum. 234 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) a slightly larger quantity of antennal sensilla placodea than Scaptotrigona, and Apis presents a larger quantity than these two. It is important to analyze the number of sensilla in relation to the size of the bee since Johnson and Howard (1987) have shown that these two characters are highly correlated. According to Kerr (1969), Nannotrigona testaceicornis has a barely evolved sys- tem of communication between workers. It has been shown that the worker that has found a food source enters the colony and produces a characteristic sound. The workers surrounding this bee immediately start to produce the same sound and in less than one minute the entire colony will be buzzing, imitating this sound, and at this time several bees leave the colony looking for food (Kerr and Esch, 1965). Thus, the bee that found the source alerts the colony in terms of the existence of this source but does not provide any indication of the location. The source is only found based on the smell of the food. In contrast, Scaptotrigona postica presents a more evolved communication system. The bee that found the food source leaves a scent trail between the source and the colony, marking flowers, leaves and branches every 1 to 2 meters with a drop of secretion (rich in compounds such as 2-heptanone, 2-nonanone and benzaldehyde) produced by the mandibular glands (Lindauer and Kerr, 1958; Kerr et al., 1963). Due to the type of communication, i.e., formation of a pheromone trail, it may be expected that Scaptotrigona postica presents a larger quantity of antennal sensilla placodea (olfactory disks) than Nannotrigona testaceicornis, which is not the case when the size of the flagellum is also considered. In this respect, the number of antennal sensilla placodea of Scaptotrigona postica, in addition to not agreeing with the type of communication they possess when compared to Apis mellifera and Me- lipona quadrifasciata (Silva de Moraes and Cruz-Landim, 1972; Stort and Barelli, 1981), also does not agree with the type of communication they possess when com- pared to Nannotrigona testaceicornis. These results support the observation that the number of antennal sensilla placodea is not simply related to the use of pheromones for the recruitment of individuals to the food source, but may play a broader role in the complex number of activities performed by bees (Johnson and Howard, 1987). ACKNOWLEDGMENTS The authors are grateful to Dr. Jose Arana Varela, Chemistry Institute, UNESP, Araraquara, for permitting the use of the scanning electron microscope, to Dr. Mario Cilense and to Mr. Sebastiao Dameto, of the same Institute, for help during the use of the equipment. The authors also wish to thank the referee for the corrections and suggestions. LITERATURE CITED Cruz-Landim, C. 1967. Estudo comparativo de algumas glandulas das abelhas (Hymenoptera, Apoidea) e respectivas implica9oes evolutivas. Arq. Zool. Sao Paulo, 15(3): 177-290. Cunha, R. A. 1973. Taxonomia numerica de alguns meliponinae (Hymenoptera — Apidae). Cienc. Biol. (Portugal) 1:25-42. Dietz, A. and W. J. Humphreys. 1971. Scanning electron microscopic studies of antennal re- ceptors of the worker honey bee, including sensilla campaniformia. Ann. Entomol. Soc. Am. 64(4) :9 19-925. Johnson, L. K. and J. J. Howard. 1987. Olfactory disc number in bees of different sizes and ways of life (Apidae: Meliponinae). J. Kans. Entomol. Soc. 60(3):380-388. 1997 ANTENNAE OF NANNOTRIGONA TESTACEICORNIS 235 Kerr, W. E. 1969. Some aspects of the evolution of social bees (Apidae). Evol. Biol. 3:119- 175. Kerr, W. E., A. Ferreira and N. S. Matos. 1963. Communication among stingless bees-additional data (Hymenoptera — Apidae). J. New York Entomol. Soc. 71:80-90. Kerr, W. E. and H. Esch. 1965. Comunica^ao entre as abelhas socials brasileiras e sua contri- bui9ao para o entendimento da sua evolu^ao. Ciencia e Cultura. 17(4):529-538. Kuwabara, M. and K. Takeda. 1956. On the hygroreceptor of the honey bee Apis mellifera. Physiol. Ecol. 7:1-6. Lacher, V. 1964. Elektrophysiologische Untersuchungen an einzelnen Rezeptoren fur Geruch, Kohlendioxyd, Luftfeuchtigkeit und Temperatur auf den Antennen der Arbeitsbiene und der Drohne. Z. Vergl. Physiol. 48:587-623. Lindauer, M. and W. E. Kerr. 1958. Die gegenseitige verstandingung bei den stachellosen Bi- enen. Z. Vergl. Physiol. 41:405-434. Lindauer, M. and W. E. Kerr. 1960. Communication between the workers of stingless bees. Bee World 41(2):29-41, 41(3):65-71. Moure, J. S. 1951. Notas sobre Meliponinae (Hym. — Apoidea). Dusenia, 2:25-70. Nogueira-Neto, P. 1970. A cria9ao de abelhas indigenas sem ferrao. Edi9ao Tecnapis - SP, 365 pp. Sakagami, S. F. and R. Zucchi. 1966. Estudo comparativo do comportamento de varias especies de abelhas sem ferrao, com especial referenda ao processo de aprovisionamento e pos- tura das cdulas. Ciencia e Cultura, 18(3):283-296. Silva de Moraes, R. L. and C. Cruz-Landim. 1972. Estudo comparativo de orgaos sensoriais em abelhas com diferentes tipos de comunica9ao. Rev. Brasil. Biol. 32(2): 185-196. Stort, A. C. 1979. Estudo de caracteres morfologicos e suas rela9oes com o comportamento de defesa de abelhas do genero Apis. Livre Docencia thesis; UNESP, Rio Claro, Brasil. 179 pp. Stort, A. C. and N. Barelli. 1981. Antennal sensory structures of Scaptotrigona postica (Hy- menoptera: Apidae). J. Kans. Entomol. Soc. 54(4):75 1-756. Stort, A. C. and M. E. Rebustini. 1997. Differences in number of sensilla coeloconica, ampul- lacea and campaniformia in the antennae of four bee groups and relationship between these sensilla and the defensive behavior of Africanized bees. J. Apic. Res. (in press). Stort, A. C. and M. M. B. Moraes-Alves. 1998. A study of the sensory structures of the antennae of Scaptotrigona postica workers (Hymenoptera — Apidae). Rev. Brasil. Biol. 58(1): 163- 167. Received 2 September 1997; accepted 20 November 1997. NOTES AND COMMENTS J. New York Entomol. Soc. 105(3-4);236-237, 1997 COCCIDULA FERRUGINEA GORHAM, A SENIOR SYNONYM OF EREMOCHILUS HOWDENI GORDON AND VANDENBERG (COLEOPTERA: COCCINELLIDAE: EPILACHINAE) Gordon and Vandenberg (1994) described Eremochilus howdeni based on two female specimens from Mexico. Roger Booth, International Institute of Entomology, London, recognized the similarity between that description and a syntype of Coc- cidula ferruginea Gorham (1888) in the Natural History Museum, London. He kindly arranged a loan of the syntype, and examination of that specimen resulted in con- firmation of the suspected synonymy. Coccidula ferruginea is here transferred to Eremochilus as a senior synonym of E. howdeni. The male syntype labeled “Type (orange bordered disc)/Toxpam/Mexico, Salle Coll.?2300(green paper)/Coccidula? ferruginea Gorh.” is designated and labeled as the lectotype of C. ferruginea. Male genitalia are illustrated in Fig. 1 and constitute the first figure published for this species. The genitalia differ in configuration from the congeneric E. weisei Gordon and Vandenberg (1987) (the only other species for which a male is available) pri- marily in possessing a tubular sipho with a much reduced capsule and an unmodified apex of the basal lobe. — Robert D. Gordon and Natalia J. Vandenberg, Systematic Fig. 1 . Eremochilus ferruginea (Gorham) (lectotype). Male genitalia: sipho (left), phallo- base and trabes from lateral and ventral views (right). 1997 NOTES AND COMMENTS 237 Entomology Laboratory, PSI, Agricultural Research Service USD A, c/o U.S. Na- tional Museum of Natural History, Washington, D.C. 20560. ACKNOWLEDGMENTS We thank Roger Booth, International Institute of Entomology, London, for alerting us to this synonymy and arranging the loan of a syntype, and Taina Litwak for creating the genitalia illustrations. For manuscript review we thank J. Chapin, Louisiana State University, Baton Rouge, and H. Dozier, Pickens, South Carolina. LITERATURE CITED Gordon, R. D. and N. J. Vandenberg. 1987. Eremochilini, a new tribe of neotropical Epilach- ninae (Coleoptera; Coccinelidae). J. New York Entomol. Soc. 95:5-9. Gordon, R. D. and N. J. Vandenberg. 1994. First record of Eremochilini (Coleoptera: Cocci- nelidae: Epilachninae) from Mexico. J. New York Entomol. Soc. 102:107-110. Gorham, H. S. 1888. Biologia Centrali-Americana, Insecta, Coleoptera, Erotylidae 7:113-128. Received and accepted 19 May 1997. J. New York Entomol. Soc. 105(3— 4):238— 242, 1997 JACQUES CARA YON (1916-1997) AN APPRECIATION With the death of Professor Jacques Carayon hemipterology has lost one of its greatest and certainly one of its most versatile students. This is not a formal obituary as I am certain that the tributes from his European colleagues will discuss more completely than I am able the many contributions of this outstanding man. Here I hope to be able to informally express my appreciation of the accomplishments of one of the outstanding figures in twentieth century entomology. Prof. Carayon unquestionably deserves a place, not only as one of the leaders in Hemipterology in this century, but as one of the leading figures in the entire history of the science. Carayon is best known for his outstanding pioneering work on “traumatic insem- ination” which he explored in a wide variety of cimicomorph taxa and demonstrated the remarkable extent of this seemingly unlikely method of copulation and sperm transmission. He discussed and illustrated this phenomenon in a series of truly ex- quisitely composed papers. For this work alone he stands in a unique position among his contemporary colleagues. However, Carayon’s interests and abilities extended far beyond the fine work on insemination mechanisms. His work on the integumentary glands is not only out- standing for its morphological and histological thoroughness, but also for its impor- tance to our understanding of higher group relationships within the Hemiptera. It certainly would be misleading to portray Jacques Carayon as primarily a mor- phologist and a field biologist. He was always a first class taxonomist and frequently his papers combined the wonderful histological work that he did with his wife with new classification schemes and often the descriptions of new taxa. Perhaps the most seminal of these papers was his large summation of the classification of the Antho- coridae. This paper proposed the first really modem schema of relationships in the family since the work of Reuter many years previously. Some Hemipterists may not be aware of the background of the subsequent series of events that have led to the present ambiguities in the classification of this group. After Carayon published his major classification of the Anthocoridae it seemed an appropriate time for a young worker to concentrate upon the taxonomy of the group. With Dr. Carayon’s approval I suggested this project to a promising young graduate student in my laboratory named Laurine Ford. Ms. Ford, who had a mastery of French, took the sum of Dr. Carayon’s work on the Anthocoridae together with all of his contributions to trau- matic insemination in related families and constmcted a phylogenetic “tree” of re- lationships. Unfortunately she left the field with this unpublished but available in thesis form. It served as part of the analysis (together with the work of Kerzhner), for a detailed cladistic analysis by Schuh & Stys in 1991 which resulted in the removal of a portion of the Anthocoridae to a separate family position. Prof. Carayon did not agree with this classification, but his accident and subsequent ill health unfortunately made it impossible for him to share with us his reasons for disagree- 1997 IN MEMORIAM 239 merit. I believe the loss of Carayon’s opinion to be a most unfortunate loss for Hemipterology. Carayon also published the first work of real consequence on several groups. Important among these was his innovative study of the peculiar family Plokiophilidae whose members live in the webs of Embioptera and spiders and which Carayon was able to show were part of the traumatic insemination complex. His work on the Pachynomidae was equally innovative and raised questions concerning the homology of trichobothria in this family relative to the Pentatomomorpha. In a similar way he treated the Nabidae as part of this complex group and made his usual original con- tributions so ably carried on by Dr. Kerzhner. In point of fact it is not an exaggeration to say that when Jacques Carayon studied a group it invariably resulted in published work that was not only remarkably complex but added new dimensions to our knowl- edge of the group both taxonomically and biologically, resulting from the morpho- logical information that was always new and provocative. His contributions alone do not tell the complete story of his contributions to Hemipterology. Of all the French workers of his generation he was perhaps the most generous of his time and in imparting his knowledge to students and to workers from other countries. He was unfailingly polite and helpful to those who visited his laboratory, spending much time with even the youngest visitors. After his mastery of English improved he was pleased to discuss scientific matters with visiting col- leagues whose lack of knowledge of French must have been frustrating to him. Carayon was perhaps first of all a naturalist, one who was always interested in the biology of the Hemiptera. To him is due our knowledge of the remarkable mating behavior in the lygaeid bugs of the genus Stilbocoris wherein the males impale small seeds upon their “beak”, partially digest them and offer them to females to enhance mating acceptance. Almost as an aside Carayon discovered during this work that the various species were ovo viviparous; to my knowledge, still the only known case of this in the Lygaeoidea. His work on this genus also unfortunately points up the many studies that he did not complete. Nearly two decades ago he told me of experiments he had conducted in his laboratory with Stilbocoris natalensis and a series of other species of Stilbocoris. In every case where both species were confined together S. natalensis would within a short period completely eliminate the other species. Both of us had hoped to follow these observations in the field, for the genus is speciose in Africa and beautifully adapted to move quickly into an area, exploit a crop of Ficus seeds, and move out. Carayon visualized a group of species that survived by adapting to getting to a seed crop quickly, producing a generation before each was overwhelmed by the ubiquitous S. natalensis. Carayon wished to revise this genus and I unfortunately dissuaded him believing another worker was already engaged. Unfortunately the revision remains to be completed. My first personal acquaintance with Prof. Carayon also involved his observations as a field naturalist. My wife and I visited his laboratory in January of 1961 during the time I was working at the then British Museum (Natural History). Carayon had discovered that a species of lygaeid of the genus Mizaldus occurred in West Africa in the nests of ploceid birds and that it fed upon beetle larvae living in the nests. This was remarkable as previously only members of one tribe in the large subfamily to which Mizaldus belonged were known to be predatory (and they fed upon ver- tebrate blood). This incidentally led to an amusing exchange between us which 240 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3-4) perhaps illustrates his character and which I have always treasured. Carayon could not identify the insect and asked if I would attempt it. I did discover that it was an undescribed species and wrote a preliminary draft of a paper describing it and dis- cussing the biological information that Prof. Carayon had given me. I sent a draft to Paris for Prof. Carayon to look over as a paper by “J. Carayon and J. Slater”. In due time it was returned with the authorship reversed. I wrote pointing out that the formal description of the new species was not the most significant part of the paper as compared to the discovery of insect feeding by a member of a tribe in which such food habits had not previously been known to occur. The answer was polite, but to the point. He, Carayon, said that he thought he was able to ascertain who had done the most significant work on the paper and who must be the senior author. There seemed no further room for discussion, I was dismissed! It was of course an act of kindness on his part, but even today I believe the authorship sequence is inaccurate. Some of my most treasured memories are of his visit to the International Congress of Entomology in Washington D.C. in 1976. I had been asked to organize a sym- posium on the status and advances in Hemipterology. It was natural to ask Jacques Carayon to be the first speaker. The symposium went well, Jacques gave it a stim- ulating start, so much so that we organized an informal discussion of relationships within the order later in the meeting. At this meeting excitement was immediate and centered around disagreements between Carayon and Rene Cobben of the Nether- lands, as I recall chiefly concerning the position of the Reduvioidea. The air was blue with the smoke from Cobben’s cigar and with the words “Jamais, jamais, c’est impossible!” that kept erupting from Carayon. I don’t think the matter was really ever settled, but it stimulated all of us. That same evening Carayon gave a more technical paper and when he arose to do so he announced that it was necessary for him to give this paper in French. His enthusiasm seemed to wane when he slowly came to realize that much of his audience was not able to follow the details of his discussion. I am convinced that he was not really aware that suddenly he began to discuss the paper in English and only occasionally lapsed into French thereafter. Thus, one remembers him not only as a scientist of the first class but also as a very human man, friendly, cooperative and a pleasure to have known personally as well as to respect professionally. Jacques Carayon was bom November 11, 1916 in Toulouse France. He was the eldest of three children. His father was a medical doctor whose premature death in 1938 together with an earlier illness of his own prevented Jacques from following his father into the medical profession. He had been interested in the Natural Sciences since he was eleven years old and when it became financially impossible for him to enter medical studies he turned to natural history and entered the University at Paris in the Natural Sciences. In 1975 Carayon fortunately summarized his career and his contributions as part of his application for the Headship of the Laboratory of Entomology at the National Museum in Paris (which he obtained). In this important summary entitled “Titres et Travaux Scientifiques de M. Jacques Carayon” he not only analyzed what he con- sidered his most important scientific work, but also fortunately one finds here the various academic and administrative positions that he held as well as his formal honors. 1997 IN MEMORIAM 241 In 1947 he married Gabrielle Carayon. This marriage was very important for his career as she was a skilled technician who became his assistant and to her must go much credit for the exquisite histological work that graced so many of his outstand- ing papers. Not all of his foreign colleagues realize that Carayon was not only a fine scientist but also from childhood until his death a devoted naturalist. He had a house in Provence where he spent long summer periods working in the field as well as on his various manuscripts and where he taught his grandchildren the joys of knowledge of the natural world. Prof. Carayon spent his entire career in Paris where he held numerous offices involving senior administrative responsibilities both at the University of Paris, the National Museum and its affiliation with the Zoological Institute of National Agron- omy. Thus throughout his career he carried an extensive administrative burden which makes his scientific contributions even more remarkable and impressive. This informal recognition of his status is not intended to discuss in detail his many offices and distinctions, but one should certainly mention his Presidency of the En- tomological Society of France as early as 1956, his expeditions to West Africa and the Cameroons in 1946 and 1947, his participation in many international entomo- logical and zoological congresses such as those in Stockholm, Amsterdam, Vienna, Moscow, Copenhagen, London, Washington, etc. culminating with his election to the Permanent Committee on International Entomological Congresses at the Kyoto Congress in 1980 (announced appropriately by his long time colleague J. C. M. Carvalho). He rose steadily through the administrative and scientific hierarchy in France and became Head of the Laboratory of Entomology of the Museum in 1975, a position which he held until his retirement in 1986. During his career he was always involved in administrative duties and he never left his obligations to eco- nomic aspects of entomology. At the time of his retirement a volume of the Annals of the Entomological Society of France was dedicated to him and included scientific articles from his colleagues around the world (this included an especially important tribute by his colleague Claude Dupuis that summarized Carayon ’s life and work in exemplary fashion. Dupuis says appropriately after lauding Carayons’ work on trau- matic insemination and on integumentary glands the following: “Lensemble, sac- compagnant, toutes les fois que necessaire, de donnees ecologiques et de taxinomie descriptive, constitue une oeuvre remarquablement coherente qui vous vaut une po- sition de leader parmi las connaisseurs du sous-ordre.”) Following his formal retirement Prof. Carayon continued to work actively, es- pecially upon the integumentary glands which he had studied for many years. In 1990 he was involved in a terrible automobile accident during a trip to Turkey which left him for a time in a coma and many months in the hospital. While he recovered mentally he really never did recover physically from this traumatic acci- dent and was unable to carry on his research program. His mind was active to the end and he never lost his interest in what his colleagues were accomplishing. There is a sadness that Carayon was not able to finish many of the important projects that he was engaged in, but we can rejoice that he was able to accomplish so much. He was a scientist of the first order of magnitude and able to do outstanding work while carrying administrative burdens and in being a stimulus and guide to many students. Hemipterology has lost one of its greatest men, a man whose accom- 242 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 105(3^) plishments will continue to be recognized and honored so long as the science con- tinues to be a subject of study. Since this is not a formal obituary I take the liberty of citing here a few of his works that I believe will give the flavor of a few of his many important contributions. LITERATURE CITED Carayon, J. 1966. Traumatic insemination and the paragenital system. In: R. L. Usinginer (ed.). Monograph of Cimicidae (Hemiptera — Heteroptera) Vol. 7, 81-166. The Thomas Say Foundation. Entomological Society of America. Lanham. Md. Carayon, J. 1970. Etude de Alloeorhynchus d’Afrique Centrale. avec quelques remarques sur la classification des Nabidae [Hemiptera]. Ann. Soc. Entomol. Er., n.s. 6:899-931. Carayon, J. 1971. Notes et documents sur I’appareil odorant metathoracique des Hemipteres. Ann. Soc. Entomol. Fr., n.s. 7:737-770. Carayon, J. 1972. Caracteres systematiques et classification des Anthocoridae (Hemipt.). Ann. Soc. Entomol. Fr., n.s. 8:309-349. Carayon, J. 1974. Etude sur les Hemipteres Plokiophilidae. Ann. Soc. Entomol. Er., n.s. 10: 409-525. Carayon, J. 1977. Insemination extra-genitale traumatique. In: R P. Grasse (ed.), Anatomie, Systematique, Biologie, Insectes, Gametogeneses, Fecondation, Metamorphoses, 351- 390. Traite de zoologie. Masson, Paris. Carayon, J. and A. Villiers. 1968. Etude sur les Hemipteres Pachynomidae. Ann. Soc. Entomol. Fr., n.s. 4:703-739. Carayon, J., R. L. Usinger and P. Wygodzinsky. 1958. Notes on the higher classification of the Reduviidae, with the description of a new tribe of the Phymatinae (Hemiptera — Heter- optera). Rev. Zool. Bot. Afr. 57:256-281. James A. Slater, Dept. Ecology & Evolutionary Biology, University of Connecticut, Storrs, CT. 1997 HONORARY MEMBERS Dr. F. S. Chew Ms. Su Zan Noguchi Swam Dr. James Forbes Dr. James A. Slater Dr. John G. Franclemont LIFE MEMBERS Dr. Annette Aiello Professor Clive R. Bailey Mr. David G. Casdorph Mr. Franklyn W. Commisso Dr. Howard E. Evans Dr. Durland Fish Mr. Irving Granek Prof. Dipl. Ing. Ernst Heiss Dr. Donald F. J. Hilton Professor Hussein S. Hussein Mr. Mark Indenbaum Dr. Kurt Johnson Dr. Gary G. Kennen Dr. J. Krikken Mr. George Ladd Dr. Brobson Lutz, M. D. Dr. Gordon A. Marsh Dr. John C. Morse Mr. M. S. Moulds Mr. Kikumaro Okano Dr. David E. Ruiter Dr. Randall T. Schuh Dr. John A. Shetterly Dr. Kathryn M. Sommerman Dr. F. Christian Thompson Mr. George F. Townes Mr. Theodore H. Weisse SUSTAINING MEMBERS Dr. Richard S. Beal, Jr. Dr. R. W. Homabrook Mr. Ralph F. Kirchner Mr. Joseph M. Laino Dr. Charles C. Porter Dr. Richard G. Robbins Mr. Enrico Sismondo REVIEWERS The editor thanks the following individuals who reviewed manuscripts published in 1997: Donat Agosti, Chuck Bellamy, Catherine N. Duckett, R. Wills Flowers, David G. Furth, Ann E. Hajek, Mary Liz Jameson, Luis F. Mendes, Edward L. Mockford, S. Nakahara, James Pakaluk, Eric Quinter, Brett C. Ratcliffe, Jerome G. Rozen, and Randall T. Schuh. CHANGE IN EDITOR Dan A. Polhemus, Department of Entomology, National Museum of Natural His- tory, MRC 105, Smithsonian Institution, Washington, DC 20560, will assume edi- torship of the Journal commencing with Volume 106. All new manuscripts should be submitted to him, using the conventions of the Instructions to Authors on the inside back cover. ( ' -'OM M •■/ tfi ;4 ; INSTRUCTIONS TO AUTHORS The Journal of the New York Entomological Society publishes original research resulting from the study of insects and related taxa. Research that contributes information on taxonomy, classification, phylogeny, biogeography, behavior, natural history, or related fields will be con- sidered for publication. The costs of publishing the Journal are paid by subscriptions, mem- bership dues, page charges, and the proceeds from an endowment established with bequests from the late C. R Alexander and Patricia Vaurie. Manuscripts should be submitted in triplicate to: Dr. James M. 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Because of limited funds, all such requests will be handled on a first-come first-served basis. Authors will receive a reprint order blank with the proofs. Reprints are ordered directly from the printer with no benefit accruing to the Society. Journal of the New York Entomological Society VOLUME 105 SUMMER-FALL 1997 NOS. 3-4 CONTENTS Morphological caste differences in the neotropical swarm-founding polistine wasps: Parachartergus smithii (Hymenoptera: Vespidae) Sidnei Mateus, Fernando Barbosa Noll and Ronaldo Zucchi Two new neotropical genera of Embiidae (Embioptera, Insecta) Claudia A Szumik A review of the genus Heliothrips (Thysanoptera; Thripidae) with a new sister- species of the greenhouse thrips from South Eastern Brazil Laurence A. Mound and Renata C. Monteiro Two new enigmatic Melophorus species (Hymenoptera: Eormicidae) from Aus- tralia Donat Agosti A new Mexican species of Homoiosternus (Coleoptera: Melolonthidae; Rutelinae) Leonardo Delgado and Julian Blackaller-Bages A new species of Metapolybia Ducke from Central America (Hymenoptera: Ves- pidae; Polistinae) Mark E. Smethurst and James M. Carpenter A new species of Loneura (Psocoptera: Ptiloneuridae) from Chiapas, Mexico Alfonso Neri Garcia Aldrete A new species of Probolomyrmex (Hymenoptera: Eormicidae) from Guanacaste, Costa Rica Sean T. O ’Keefe and Donat Agosti New species and a new name for Antillean Buprestidae (Coleoptera) Henry A. Hespenheide The ant (Hymenoptera: Eormicidae) types in Argentinian collections Donat Agosti Notes on the genus Thasus (Hemiptera: Coreidae) Carl W. Schaefer and Richard J. Packauskas The real identity of Discoelius strigosus costarricensis Bertoni (Hym.: Vespidae: Eumeninae) Bolivar R. Garcete Barrett Observations on phenology, development, and mortality of larvae of the hazelnut weevil (Curculio obtiisus (Blanchard): Curculionidae) in nuts of beaked hazelnut {Corylus cornuta Marshall: Betulaceae) in thickets in Maine L. W. Treadwell and R. H. Storch Sensory structures of the antennae of Nannotrigona testaceicornis (Apidae: Me- liponinae) Antonio Carlos Stort and Monica M. B. Moraes-Alves Notes and Comments Coccidula ferruginea Gorham, a senior synonym of Eremochilus howdeni Gordon and Vandenberg (Coleoptera: Coccinellidae: Epilachinae) Robert D. Gordon and Natalia J. Vandenberg In Memoriam Honorary, Life, and Sustaining Members Reviewers for 1997 129-139 140-153 154-160 161-169 170-179 180-185 186-189 190-192 193-198 199-205 206-214 215-220 221-229 230-235 236-237 238-242 243 243 Change in Editor 243 HECKMAN BINDERY INC. FEB 99 -To-Pleas#’ N. MANCHESTET INDIANA 46962